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OSMANIA  UNIVERSITY  LIBRARY 

Call  No.  Sl7.$/(Sf4f/          Accession  No.    3 
Author 


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HANDBOOKS  OF  AMERICAN  NATURAL  HISTORY 
ALBERT   HAZEN   WRIGHT,   ADVISORY    EDITOR 


Handbook  of  Frogs  and  Toads 

ALBERT  HAZEN  WRIGHT  AND  ANNA  ALLEN  WRIGHT 


HANDBOOK  OF 

Frogs  and  Toads 

OF  THE  UNITED  STATES  AND  CANADA 

BY  ALBERT  HAZEN  WRIGHT 

PROFESSOR    OF    ZOOLOGY,    EMERITUS,    CORNELL   UNIVERSITY 

AND  ANNA  ALLEN  WRIGHT 


THIRD        EDITION 


Comstock  Publishing  Company,  Inc. 

ITHACA,  .NEW  YORK,  1949 


COPYRIGHT  1933,  1942,  1949  BY 
COMSTOCK  PUBLISHING  COMPANY,  INC. 


All  rights  reserved.  This  book,  or  parts  thereof,  must 
not  be  reproduced  in  any  form  without  permission  in 
writing  from  the  publisher,  except  by  a  reviewer  who 
wishes  to  quote  brief  passages  in  a  review  of  the  book. 


PRINTED  IN  THE  UNITED  STATES  OF  AMERICA  BY  THE 
VAIL-BALLOU    PRESS,    INC.,    BINGHAMTON,    NEW    YORK 


This  volume,  meant  to  serve  the  public,  the  scientist, 
and  the  group  treated,  is  dedicated  to  the  four  Amer- 
ican women  who,  in  addition  to  serving  the  public 
and  science  generously,  have  in  the  last  half-century 
contributed  most  notably  to  the  study  of  this  group: 

Mary  Hewes  Hinckley,  1845-1944 
Mary  Cynthia  Dickerson,  1866-1923 
Helen  Dean  King,  1869- 
Helen  Thompson  Gaige,  1889- 


Preface 


NINETY  YEARS  ago  John  Le  Conte  in  the  prefatory  paragraphs  to 
his  "Descriptive  Catalogue  of  the  Ranina  of  the  United  States"  (Gen., 
1856,  p.  423)  wrote: 

Before  I  begin,  it  is  necessary  to  observe  that  all  the  Ranina  which  I  have  ever 
seen  have  more  or  less  the  power  of  changing  color  at  will.  The  character  of 
color,  therefore,  of  so  much  moment  in  the  description  of  many  other  animals, 
is  here  of  very  little  value;  for  none  of  the  marks  dependent  on  it  are  constant.  In 
consequence,  it  requires  numerous  specimens,  living  subjects  and  long  study  to 
produce  any  description  that  approaches  perfection. 

How  far  I  have  succeeded  in  my  attempt  remains  to  be  seen.  I  have  been  long 
and  sedulously  engaged  in  my  researches.  Every  description  has  been  made  from 
living  specimens. 

In  conclusion  (p.  431)  he  observed: 

These  are  all  the  species  of  this  family  of  reptiles  which  I  have  been  able  to 
see  in  a  living  state.  I  hold  it  to  be  impossible  for  any  one  to  make  a  correct  de- 
scription of  an  animal  which  has  the  power  of  changing  its  color  at  will,  unless 
he  has  it  alive;  and  this  power  they  all  possess  in  a  greater  or  less  degree;  when 
preserved  in  alcohol  they  fade.  The  difficulty  of  procuring  descriptions  made  on 
the  spot  where  the  objects  are  found,  has  rendered  this  part  of  natural  history  as 
confused  and  unsatisfactory  as  the  researches  of  botanists  who  draw  up  their 
accounts  of  plants  from  dried  specimens. 

In  all  our  portrayals  we  have  sought  to  emphasize  the  living  animal  by 
photographs  from  life,  by  color  descriptions  from  life,  by  journal  notes  from 
the  field,  and  particularly  by  excerpts  from  others'  work  pertaining  to  live 
frogs.  In  many  regions  we  have  been  transitory  visitors  and  the  resident 
naturalists  can  tell  a  more  complete  story.  Our  notes  and  the  notes  of 
others,  which  in  the  two  earlier  editions  were  merged  under  the  heading 
"Notes,"  are  here  expanded  and  designated  "Journal  notes"  and  "Authorities' 


corner." 


We  have  the  warmest  feelings  for  the  countless  friends  and  strangers  who 
have  helped  us  at  different  times  and  in  different  places.  We  have  seen  most  of 
the  preserved  materials  of  the  important  collections  and  we  thank  their  cus- 
todians. An  especial  effort  has  been  made  to  secure  live  material  and  many 


viii  PREFACE 

people  have  contributed.  The  first  group  constitutes  individuals  at  most  of  the 
large  zoos  of  the  country.  These  people  offered  to  loan  or  in  some  instances 
gave  us  much  needed  material.  In  a  similar  spirit  specialists  or  workers — one 
hundred  or  more — in  many  places  and  institutions  served  us  in  countless 
fashions.  The  supply  houses  and  biological  bureaus  were  very  eager  and  will- 
ing to  help.  The  list  qf  old  students  who  have  mailed  or  shipped  us  live  frogs 
would  be  hard  to  compile.  Many  have  contributed  to  these  almost  weekly 
surprises  through  many  years.  The  limits  of  this  book  forbid  such  a  list. 
When  we  recall  that,  in  California  colleges  alone,  we  have  some  twelve  to 
fifteen  teachers  of  biology,  old  students,  we  realize  the  size  of  our  obliga- 
tions. Throughout  this  book  will  appear  the  collectors,  helpers,  comrades — 
our  creditors. 

For  the  typing  of  this  work  we  extend  our  thanks  to  Mrs.  Katharine  Kapp, 
Dr.  Ann  L.  Dunham,  Mrs.  Frankie  Culpepper  Goerges,  and  Mrs.  R.  F.  Darsie. 
Two  associates  who  have  helped  us  particularly  are  Professors  W.  J.  Hamilton, 
Jr.,  and  Edward  C.  Raney.  We  are  also  indebted  to  Cornell  University  for  the 
Faculty  Research  Grant  to  help  in  typing,  purchase  of  specimens,  visits  to 
museums,  and  trips  for  specimens. 

ALBERT  HAZEN  WRIGHT 
ANNA  ALLEN  WRIGHT 
Ithaca,  New 
December, 


Contents 


PREFACE  .  ...  .    vii 

GENERAL  ACCOUNT      .  .  .  .     .          .  i 

Common  names  .  i 

Present  classifications  .  ....  .  2 

Scientific  name  .  .2 

Specific  evaluation  of  our  North  American  forms  .  4 

Range  5 

Habitat  .  ....  .10 

Size     .  .  .     .  ...          .          10 

General  appearance  .  .  12 

Color  .     .  .  ...  .12 

Structure  .          .          ....     12 

Voice 12 

Vocal  sacs  .          .  .  ...     14 

Color  of  throat  .  .          .     .     .     18 

Enlarged  tympana  (males)  .  .  20 

Breeding  sizes  .     20 

Summary  of  secondary  sexual  characters  .  .     24 

Ovulation       .  .26 

Egg-laying  process  .  .     31 

Eggs  .          ...     31 

Synopsis  of  eggs  of  United  States  frogs  .  .     36 

Tadpoles        .  .  .46 

Synopsis  of  United  States  tadpoles  .  .          .     .     .*    54 

Development  and  transformation  .  .  75 

Journal  notes  .  ....     79 

Authorities'  corner        .  ...  -79 

KEYS  .  .  81 

Families  .          .  -          -  .     .  .     .     .     81 

Ascaphidae  ...  .  .     .     .     .     81 

Scaphiopodidae  ...  82 

Bufonidae  83 

Hylidae      .  .          ...  87 

Leptodactylidae  .     .  ...          .     .          94 

Ranidae  ...  ...     96 

Brevicipitidae  .  .  103 


x  CONTENTS 

Page  Number  of 

Account  Plate  Map 

ACCOUNTS  OF  SPECIES  104 

Ribbed  toads,  Ascaphidae  104 

American  bell  toad,  Ascaphus  truei  105  no  105 

Spadefoots,  Scaphiopodidae  in 

Couch's  spadefoot,  Scaphiopus  couchii  in  no  112 

Hammond's  spadefoot,  Scaphiopus  hammondn  hammondii      113  117  114 

Central  Plains  spadefoot,  Scaphiopus  hammondii  bombifrons  116  117  114 
Great  Basin  spadefoot  toad,  Scaphiopus  hammondii  inter- 

montanus  120  120  114 

Spadefoot,  Scaphiopus  holbroofyi  holbroo1(ii  123  125  124 

Key  West  spadefoot,  Scaphiopus  holbroo\n  albus  127  128  124 

Hurter's  spadefoot,  Scaphiopus  holbrootyi  hurterii  130  130  124 

Toads,   Bufonidae  135 

Colorado  River  toad,  Bufo  aharius  135  139  136 

American  toad,  Bufo  amcricanus  americanus  140  139  144 

Hudson  Bay  toad,  Bufo  americanus  copei  143  147  144 

Northwestern  toad,  Bufo  boreas  boreas  146  147  144 

California  toad,  Bufo  boreas  halophilus  150  153  151 

Amargosa  toad,  Bufo  boreas  nelsoni  154  153  151 

Southern  California  toad,  Bufo  californicus  156  161  136 

Yosemite  toad,  Bufo  canorus  160  161  151 

Great  Plains  toad,  Bufo  cognatus  .   164  168  144 

Spade-footed  toad,  Bufo  compactihs  167  168  136 

Little  green  toad,  Bufo  debilis  172  175  172 

Black  toad,  Bufo  exsul  176  175  151 

Canadian  toad,  Bufo  hemiophrys  180  183  144 

Sonoran  toad,  Bufo  insidior  184  183  172 

Marine  toad,  Bufo  marinus  188  193  144 

•Spotted  toad,  Bufo  punctatus  192  193  195 

Oak  toad,  Bufo  quercicus  197  200  207 

Southern  toad,  Bufo  terrestris  199  200  140 

Nebulous  toad,  Bufo  valliccps  203  205  195 

Rocky  Mountain  toad,  Bufo  woodhoush  woodhousii  206  205  207 

Fowler's  toad,  Bufo  woodhousii  fowleri  210  215  207 

Tree  frogs,  Hylidae  217 

Cricket  frogs,  Acris  217 

Cricket  frog,  Acris  gryllus  gryllus  217  221  218 

Cricket  frog,  Acris  gryllus  crepitans  220  221  218 

Chorus  frogs,  Pseudacris  229 

Mountain  chorus  frog,  Pseudacris  brachyphona  229  233  230 

Brimley's  chorus  frog,  Pseudacris  brimleyi  234  233  230 


CONTENTS  xi 

Account  Plate  Map 

Swamp  cricket  frog,  Pseudacns  nigrtta  mgrtta  236  239  237 

Clarke's  chorus  frog,  Pscudacns  ntgnta  clartyi  240  239  267 

Eastern  chorus  frog,  Pseudacns  mgnta  jenarum  244  249  237 

Northern  striped  tree  trog,  Pseudacns  mgrita  septentrionahs  250  249  267 

Striped  tree  frog,  Pseudacns  mgrita  tnsenata  253  259  267 

Florida  chorus  frog,  Pseudacns  mgnta  verrucosa  258  259  237 

Western  chorus  frog,  Pseudacns  occtdentalts  262  265 

Little  chorus  frog,  Pseudacns  ocular js  264  265  267 

Ornate  chorus  frog,  Pseudacns  ornata  268  273  230 

Strecker's  ornate  chorus  frog,  Pseudacns  strecf^en  272  273  230 

Tree  frogs,  Hyla  281 

Anderson  tree  frog,  Hyla  andersonn  281  284  282 

Canyon  tree  trog,  Hyla  aremcolor  287  289  293 

Viosca's  tree  frog,  Hyla  avivoca  294  297  295 

Mexican  tree  frog,  Hyla  baud  inn  298  297  293 

Creen  tree  frog,  Hyla  cineiea  cineiea  303  307  303 

Miller's  tree  irog,  Hyla  cinerea  evittata  308  307  303 

Peeper,  Hyla  c  rucijet  crucifer  311  316  312 

Florida  peeper,  Hyla  oucijer  bartramiana  315  316  312 

Pine  woods  tree  Irog,  Hyla  fetnoralis  319  325  295 

Barker,   Hyla  giattosa  324  325  282 

Pacific  tree  frog,  Hyla  regilla  329  337  267 

Giant  tree  frog,  Hyla  septentrionahs  338  337  282 

Squirrel  tree  frog,  Hyla  squirella  341  345  342 

Common  tree  toad,  Hyla  vcrsicolor  versicolor  344  345  293 

Cope's  tree  frog,  Hyla  versicolor  chrysoscelis  347  348  293 

Dusky  tree  toad,  Hyla  veisicolor  phaeocrypta  353  354 

Sonora  tree  frog,  Hyla  wnghtorum  360  348  267 

Robber  frogs,  Leptodactyhdae  366 

Mexican  cliff  trog,  Eleutherodactylus  augusti  366  369 

Texan  cliff  frog,  Eleuthei  odactylus  latrans  368  375  371 

Ricord's  frog,  Eleutherodactylus  ncordn  plamrostns  377  369  371 

White-lipped  frog,  Leptodactylus  labtalis  383  385  371 

Camp's  frog,  Syrrhophus  cam  pi  389  391  371 

Gaige's  frog,  Syrrhophus  gaigeae  394  371 

Marnock's  frog,  Syrrhophus  marnocfyi  395  391  371 

Frogs,  Ranidae  .  399 

Texas  gopher  frog,  Rana  areolata  areolata  399  403  404 

Northern  gopher  frog,  Rana  areolata  circulosa  402  403  404 

Oregon  red-legged  frog,  Rana  aurora  aurora  410  415  411 

California  red-legged  frog,  Rana  aurora  drayton'n  416  415  411 

California  yellow-legged  frog,  Rana  boyht  boylii  419  425  420 


xii  CONTENTS 

Account  Plate  Map 

Sierra  Madre  yellow-legged  frog,  Rana  boylii  muscosa  .  424  425  420 

Sierra  Nevada  yellow-legged  frog,  Rana  boylii  sierrae     .  429  433  420 

Florida  gopher  frog,  Rana  capito                                      .  434  433  404 

Slater's  frog,  Rana  cascadae                                    .           .  439  443  411 

Bullfrog,  Rana  catesbeiana                           .      .           ...  444  443  449 

Green  frog,  Rana  clamitans                                               .  450  453  451 

Nevada  frog,  Rana  fishcri  .  454  507  499 

Southern  bullfrog,  Rana  grylio                                .      .  .  459  466  460 

River-swamp  frog,  Rana  hecl(schen                          .           .  .  465  466  467 

Utah  frog,  Rana  onca  .  473  477  499 

Pickerel  frog,  Runa  palustris                                    ....  476  477  467 

Meadow  frog,  Rana  pipiens  pipiens                              .      .  .481  484  498 

Plain  meadow  frog,  I  Mutant  J  Rana  pipiens  burnsi  483  484  499 

Mottled  meadow  frog,  [Phase]  Rana  pipiens  \andiyohi  488  493  499 

Southern  meadow  frog,  Rana  pipiens  sphenocephala        .  .  492  493  499 

Western  spotted  frog,  Rana  pretiosa  pretiosa  520  528  451 

Nevada  spotted  frog,  Rana  pretiosa  luteiventns  527  528  521 

Mink  frog,  Rana  septentrionalis  .  533  534  451 

Dusky  gopher  frog,  Rana  sevosa        .                 .                .  537  404 

Wood  frog,  Rana  sylvatica  sylvatica  540  534  541 

Northern  wood  frog,  Rana  sylvatica  cantabrigensis  544  545  541 

Mexican  frog,  Rana  tarahumarae  553  557  521 

Sphagnum  frog,  Rana  uirgatipes                              .  556  557  460 

Narrow-mouth  toads,  Brevicipitidae                                  .      .  561 

Taylor's  toad,  Hypopachus  cuneus  561  563  562 

Mitchell's  narrow-mouth  toad,  Microhyla  areolata  568  562 

Narrow-mouth  toad,  Microhyla  carolinensis  571  579  562 

Taylor's  microhyla,  Microhyla  mazatlanensis  .   574  562 

Texas  narrow-mouth  toad,  Microhyla  olivacca  .  578  579  562 

BIBLIOGRAPHY  584 

INDEX                           .                     ...  623 


Handbook  of  Frogs  and  Toads 


General  Account 


TN  THE  TREATMENT  of  each  species,  the  topical  outline  is  as  follows: 
JL  Common  names,  Scientific  name,  Range,  Habitat,  Size,  General  appear- 
ance, Color,  Structure,  Voice,  Breeding,  Journal  notes,  and  Authorities'  corner. 

Common  names:  We  of  the  United  States  and  Canada  have  concerned  our- 
selves little  with  distinctive  names  for  the  amphibians.  Most  of  the  common 
names  of  the  salamanders  and  frogs  are  collective.  Just  as  all  salamanders  are 
generally  called  water  dogs  or  lizards,  so  also  the  frogs  are  known  by  a  few 
common  names  (toads,  frogs,  or  tree  frogs).  Some  people  use  the  word 
"peeper"  indiscriminately  for  several  small  frogs  that  call  in  the  early  spring. 
Others  call  peepers  lizards. 

Normally  we  expect  common  names  to  come  from  the  people  at  large,  but 
with  amphibians  and  reptiles  most  of  the  common  names  in  literature  are 
really  bookish  names.  Many  are  translations  of  the  scientific  names.  The  names 
may  come  from  widely  different  sources,  of  which  the  following  arc  a  few: 

(1)  The  person  after  whom  the  species  is  named.  Example:  Couch's  spade- 
foot.  Scaphiopus  cotichii  Baird. 

(2)  The  person  who  named  the  species.  Example:  Viosca's  tree  frog,  Hyla 
avivoca  Viosca. 

(3)  The  person  who  first  collected  it.  Example:  Taylor's  toad.  Hypopachus 
cuneus  Cope. 

(4)  A  country.  Examples:  Canadian  toad,  American  bell  toad,  Mexican 
toad. 

(5)  A  state  or  province.  Examples:  Sonora  hyla,  Winnipeg  toad,  California 
red-legged  frog. 

(6)  Habitats.  Examples:  River-swamp  frog,  pond  frog,  house  frog,  salt- 
marsh  frog,  canyon  tree  toad,  desert  tree  toad,  crayfish  frog,  gopher  frog,  wood 
frog,  savanna  cricket,  cliff  frog. 

(7)  Habits.  Examples:  Chameleon  tree  frog,  solitary  spadefoot,  grasshop- 
per frog. 

(8)  Structural  characters.  Examples:  Ribbed  toad,  narrow-mouthed  toad, 
toothless  frog,  femoral  hyla,  thick-skinned  frog. 

(9)  Voice.  Examples:  Bell  frog,  screaming  frog,  pig  frog,  rattler,  chorus 
frog,  cricket  frog. 

(10)  Color.  Examples:  Three-lined  tree  frog,  striped  tree  frog,  ornate  tree 
frog,  green  toad,  cinereous  hyla. 

(n)  Seasons.  Examples:  Spring  peeper,  shad  frog. 


2  HANDBOOK  OF  FROGS  AND  TOADS 

(12)  Miscellaneous  sources  of  many  kinds.  Folklore:  example— charming 
toad  (legend  says  this  toad,  Bufo  terrestris,  turns  your  eye  green).  Use  (bait) : 
example— pickerel  frog.  Weather  signs:  example— rain  frog.  Odor:  example- 
mink  frog. 

Present  classifications:  This  work  has  primarily  the  purpose  of  presenting 
the  living  animal.  Through  our  forty  years  of  study  and  teaching  we  have 
realized  the  inadequacy  of  vertebrae,  skull,  pectoral  girdle,  sacral  diapophyses, 
teeth,  disks,  and  other  structural  characters  (largely  osteological)  in  the  classi- 
fication of  a  smooth  (scaleless)  group.  We  early  sought  to  work  out  our  North 
American  life  histories  to  supplement  this  lack  but  doubt  if  we  will  ever  have 
knowledge  enough  to  undertake  the  task  or  to  clutter  up  scientific  literature 
with  preliminary  suggestions.  One  of  the  serious  attempts  in  this  country  has 
been  Dr.  CJ.  K.  Noble's  doctoral  dissertation,  The  Phytogeny  of  the  Salientia, 
wherein  he  attempts  to  employ  musculature  as  a  supplement  to  osteology,  as 
has  been  done  for  birds  and  mammals.  And  his  study  will  surely  have  to  be 
considered  in  the  final  analysis.  But  many  forms  need  to  be  canvassed  and 
many  life  studies  undertaken  before  we  rely  too  implicitly  on  any  set  of  mor- 
phological characters.  In  the  past  many  morphological  osteological  criteria 
have  been  overworked  and  overemphasized,  and  we  are  in  a  state  of  judicial 
suspense  awaiting  a  nonhasty  synthesis.  Therefore  we  have  temporized  by 
employing  Stejneger  and  Harbour's  check  list  with  no  particular  emphasis  of 
our  own  devising. 

Scientific  name:  Any  consideration  of  the  scientific  name  that  an  animal 
bears  implies  an  understanding  of  the  scheme  of  classification.  All  living 
things  fall  into  two  groups  or  kingdoms.  The  plants  are  treated  in  the  science 
of  botany,  the  animals  in  zoology.  The  animal  kingdom  has  several  major  sub- 
divisions or  phyla,  the  last  being  the  Vertebrata  (vertebrates).  In  the  vertebrate 
phylum,  the  various  classes  are  known  as  fishes  (Pisces),  amphibians  (Am- 
phibia), reptiles  (Reptilia),  birds  (At/es)^  and  mammals  (Mammalia).  We 
designate  the  study  of  fishes  as  ichthyology,  that  of  birds,  ornithology,  that 
of  mammals,  mammalogy,  but  we  group  together  amphibians  and  reptiles  in 
the  science  of  herpctology.  This  merging  of  the  two  groups  is  in  a  measure 
due  to  our  inability  to  designate  infallible  characters  of  separation.  A  fish  has 
fins,  a  bird,  feathers,  a  mammal,  hair,  but  reptiles  and  amphibians  have  no  one 
positively  distinctive  character. 

The  amphibians,  to  which  the  order  of  frogs  (Salientia  or  Ecaudata)  be- 
longs, have  been  variously  defined.  Some  fifteen  to  twenty  characters  have 
been  employed.  Most  living  amphibians  have  naked  skin  and  a  larval  aquatic 
stage.  Normally,  as  tadpoles  or  larvae,  they  breathe  with  gills  air  dissolved  in 
the  water;  and  as  adults  they  breathe  with  lungs.  Two  of  the  membranes  about 
a  developing  mammal  are  absent  in  amphibians.  There  are  three  living  orders: 

Apoda  (caecilians)  are  limbless,  blind,  and  wormlike.  None  occur  in  the 
United  States  or  Canada. 


GENERAL  ACCOUNT  3 

Caudata  (salamanders)  arc,  as  adults,  tailed. 

Salientia  (frogs)  are,  as  adults,  tailless. 

Seven  families  of  Salientia  or  ecaudate  amphibians  are  represented  by  99 
species  or  subspecies  in  the  United  States  and  Canada.  Family  names  in  zool- 
ogy and  botany  end  in  idae.  These  seven  families  with  the  number  of  species 
and  subspecies  in  the  United  States  and  Canada  are: 
/)  Bell  toads,  Ascaphidae  i  species 

2)  Spadefoots,  Scaphiopodidae  7  species  and  subspecies 

j)  Toads,  Bufonidae  21  species  and  subspecies 

4)  Tree  toads,  Hylidae  30  species  and  subspecies 

5)  Robber  frogs,  Leptodactylidae  7  species 

6)  Frogs,  Ranidae  28  species,  subspecies  and  phases 

7)  Narrow-mouthed  toads,  Brevici-      5  species 

pitidae 

Some  of  these  families  are  divided  into  subfamilies.  The  ending  for  sub- 
family names  is  inae.  Thus,  the  true  frogs  considered  in  this  work  belong  to 
the  family  Ranidae  and  the  subfamily  Raninae. 

The  family  is  divided  into  genera  and  the  genera  into  species. 

Ordinarily  a  scientific  name  consists  of  three  parts:  the  first  name  is  the 
generic  name,  the  second  the  specific  name,  and  the  third  the  author  or  de- 
scriber  who  first  gave  the  name.  The  specific  Latin  name  serves  as  an  adjective 
agreeing  in  gender  and  number  with  the  generic  name,  which  is  treated  as  a 
noun.  This  is  the  binomial  system  of  nomenclature.  The  meadow  frog  might 
serve  as  an  example.  It  is  called  Rana  pipiens  Schreber.  The  generic  name  is 
written  with  a  capital  and  the  specific  name  with  a  small  letter.  If  the  species 
be  divided  into  one  or  more  subspecies  or  races,  the  name  may  consist  of  four 
parts,  namely,  genus,  species,  subspecies,  and  authority.  Such  a  name  is  an 
example  of  trinomial  nomenclature.  The  swamp  cricket  frogs,  Pseudacris 
nigrita  (Le  Conte),  may  be  divided  into  several  subspecies  written  thus,  Pseu- 
dacris  nigrita  nigrita  (Le  Conte);  P.  n.  septentrionalis  (Boulenger),  etc. 
Notice  that  abbreviations  may  be  employed  after  Pseudacris  and  nigrita  have 
been  spelled  out  once. 

CATEGORIES 

Kingdom Animal  Subfamily     Raninae 

Phylum    Vertebrata  Genus  Rana 

Class  Amphibia  Species  boylii 

Order   Salientia  Subspecies  sierrae 

Family  Ranidae  Authority    Camp 

Name:  Rana  boylii  sierrae  Camp 

[Note:  Observe  that  we  use  in  this  edition  another  example,  Rana  boylii 
sierrae  Camp.  In  the  1942  edition,  we  used  Rana  pipiens  burnsi  (Weed)  with 


4  HANDBOOK  OF  FROGS  AND  TOADS 

a  purpose.  Never  have  we  been  queried  by  youthful  or  elder  scholars  on  any 
other  issue  as  on  this  example.  We  employed  it  to  stimulate  work;  unlike 
most  of  the  critics,  we  saw  these  frogs  in  the  field.  Never  did  we  seriously  hold 
it  to  be  a  valid  form,  but  in  a  book  of  this  sort  evidence  ought  to  be  presented 
with  not  too  much  youthful  certainty  or  elderly  obstinacy.] 

Specific  evaluation  of  our  North  American  forms.  About  70  of  our  99  spe- 
cies or  subspecies  are,  in  our  opinion,  established  on  firm  grounds.  If  all  the 
recently  described  forms  were  as  certain  as  Ascaphus  truei  Stejneger,  Rana  vir- 
gatipes  Cope,  R.  septentrionalis  Baird,  R.  catesbeiana  Shaw,  and  R.  hec1(scheri 
Wright,  no  hesitation  would  be  manifest. 

We  have  included  accounts  of  all  the  forms  of  Stejneger  and  Harbour's 
check  list.  The  last  species,  R.  hecl(scheri,  we  never  would  have  dared  to  de- 
scribe from  preserved  specimens  alone  had  we  not  first  seen  its  queer  tadpoles 
and  heard  its  peculiar  voice.  Nevertheless,  we  frankly  put  it  in  the  list  of  forms 
for  which  more  evidence  is  needed.  In  other  words,  we  need  field  studies  and 
much  more  ample  material  on  at  least  29  forms.  They  may  all  be  tenable, 
but  calm  judgment  dictates  that  material  alive  and  preserved,  life  histories, 
and  other  evidence  be  at  hand  before  they  rank  with  established  forms  like 
A.  truei,  Scaphiopus  couchii,  Bufo  debilis,  Hyla  gratiosa,  and  R.  virgatipes. 
The  29  forms  that  need  more  attention  are  as  follows: 

/)  Scaphiopus  hammondii  bombtfrons 

2)  Scaphiopus  hammondii  intermontanus 

j)  Scaphiopus  holbrool(ii  albtts 

4)  Bufo  boreas  nelsoni 

5)  Bufo  calif ornicus 

6)  Bufoexsul 

7)  Bufo  compactihs  (of  Southwest) 

8)  Bufo  hemiophrys 

9)  Bufoinsidior 
w)  Acris  crepitans 

//)  Pseudacris  nigrita  (6  subspecies) 

12)  Pseudacris  occidentalis  (probably  nonexistent) 

/^)  Hyla  cinerea  evittata 

14)  Hyla  crucifer  bartramiana 

75)  Hyla  wrightorum 

16)  Hyla  versicolor  chrysoscelis 

77)  Hyla  versicolor  phaeocrypta  (avivoca  may  be  the  same) 

/£)  Eleutherodactylus  augusti 

79)  Syrrhophus  campi 

20)  Syrrhophus  gaigeae 

21)  Rana  areolata  (subspecies) 

22)  Rana  boylii  (subspecies) 
Ranacascadae 


GENERAL  ACCOUNT  5 

24)  Rana  pipiens  burnsi  (color  phase) 

25)  Rana  pipicns  tyndiyohi  (color  phase) 

26)  Rana  pipiens  (subspecies  of  the  check  list) 

27)  Rana  sylvatica  latiremis 

28)  Microhyla  areolata 

29)  Microhyla  mazatlanensis 

The  99  species  and  subspecies  have  been  established  on  different  criteria 
and  degrees  of  variations.  It  is  the  age-old  problem.  What  is  a  species  or  sub- 
species? Are  similarities  in  measurements  indicative  of  relationship  or  of  the 
same  environment  (e.g.,  short  mouths  and  short  legs  in  northern  forms  in 
contrast  to  longer  mouths  and  hind  legs  in  southern  forms)  ?  Are  rugosities 
(Pseudacris,  Acris,  Microhyla,  Scdphiopus)  sufficient  for  species  or  subspecific 
distinctions?  Are  they  inherent  or  are  they  peculiar  to  certain  southern  or 
southwestern  areas  ? 

Is  the  absence  of  spots  (R.  p.  burnsi)  in  a  form  derivative  from  a  spotted 
form  (jR.  pipiens)  sufficient  for  a  species,  or  is  it  a  mutant,  a  recessive,  or  a 
metachroistic  phase  ?  We  were  of  the  group  who  had  doubts  regarding  R.  p. 
burnsi  and  R.  p.  \andiyohi,  yet  they  were  two  types  found  among  thousands 
of  live  frogs.  Had  they  come  as  a  few  specimens  (preserved  material)  in  a 
collection  from  distant  countries,  we  might  with  less  knowledge  have  de- 
scribed them  and  accepted  them  readily. 

The  29  forms  listed  above  are  some  of  the  most  interesting  and  engaging 
ones  for  future  study,  and  extended  comments  upon  each  appear  under  the 
species  accounts. 

Range:  Maps  i  to  3  give  life  zones  of  geographical  distribution  and  rainfall. 
Maps  4  to  37  show  species  distribution.  The  core  of  this  information  will  be 
found  in  A  Chec\  IJst  of  North  American  Amphibians  and  Reptiles  by  Leon- 
hard  Stejneger  and  Thomas  Barbour  (Cambridge:  Harvard  University  Press, 
1943).  We  have  added  our  own  records  through  the  years  and  such  records  as 
we  have  found  in  some  of  the  smaller  collections. 

The  ranges  were  compiled  from  dot  maps  of  all  records  for  each  form.  They 
outline  the  outer  bounds  of  each  form  but  can  in  no  way  give  all  ecological 
evaluations  each  critic  might  wish.  They  are  meant  to  give  the  layman  a  mac- 
roscopic view  of  the  range  of  a  species  or  subspecies  without  portraying  indi- 
vidual dot  records.  Sometimes  the  map  unavoidably  includes  a  state  in  which 
there  is  no  published  or  known  record.  Our  ranges  whenever  they  enter  Mex- 
ico must  be  interpreted  as  approximate. 

Geologic  speculations:  Many  features  must  be  weighed  in  a  study  of  distri- 
bution. Tolerance  of  drought  and  of  high  or  low  temperatures,  as  well  as  the 
type  of  food  and  shelter  available,  must  be  considered.  The  respective  ranges 
of  tolerance  of  concentrations  of  salt,  sulphur,  borax,  potash,  or  gypsum  must 
be  noted.  The  susceptibility  of  either  tadpoles  or  adults  may  limit  the  range. 
Frogs  are  probably  very  responsive  to  the  rock  structure  of  their  abodes,  their 


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GENERAL  ACCOUNT  7 

breeding  spots,  at  least,  being  dependent  upon  the  surface  water  of  the  region. 

Do  Pscudacris  (cricket  frogs)  dislike  granitic  and  other  igneous  situations? 
There  is  a  strange  lack  of  records  of  these  small  frogs  in  eastern  Canada  on  the 
crystalline  Laurentian  Shield,  although  one  form  extends  above  the  Arctic 
Circle  west  of  Hudson  Bay.  The  widespread  Pseudacris  n.  triscriata  (Map  18) 
stops  short  at  the  lava  plains  of  the  Snake  River  in  Idaho  but  extends  a  knob 
of  occurrence  into  the  corner  of  that  state  near  Montana  where  Carboniferous 
and  Algonkian  rocks  occur  and  into  northeastern  Utah  where  there  is  lime- 
stone. Look  at  the  range  of  P.  brachyphona  (Map  16).  Those  frogs  seem  to 
enjoy  the  carboniferous  Pennsylvanian  rocks  of  western  Pennsylvania,  West 
Virginia,  corners  of  Ohio  and  Kentucky — the  Allegheny  Plateau.  P.  n.  fen- 
arum  (Map  17)  of  Carlisle,  Pa.,  occurs  in  the  vicinity  of  Brunswick  shale  and 
other  shales  and  limestones.  Do  marine  limestone  and  marine  shell  marl  help 
produce  the  rougher  skin  otP.n.  verrucosa  (Map  17)  ?  Do  those  same  marine 
limestones  of  Key  West  influence  the  color  of  Scaphiopus  h.  albus  (Map  7), 
vary  the  Microhyla  species  (Map  37)  of  the  region,  or  affect  the  ventral  color 
of  Rana  pipiens? 

Look  for  a  moment  at  Rana  p.  prctiosa  and  12.  p.  luteiventris  (Map  35).  The 
latter  tolerates  the  tertiary  volcanic  rocks  of  the  Columbia  River  Plateau  and 
ranges  through  the  lava-infested  Snake  River  Valley  and  its  tributaries  into 
northern  Nevada,  while  R.  p.  prctiosa  prefers  the  continental  deposits  in  and 
west  of  the  Willamette  Valley  of  Oregon,  crossing  Washington  and  coming 
down,  behind  the  12.  p.  luteiventris  range,  through  the  area  of  the  Bitterroot 
Mountains  and  Grand  Tetons  to  the  Wasatch  formations  of  northeastern 
Utah. 

Does  Hyla  c.  evittata  live  at  the  fringes  of  the  range  of  H.  c.  cinerea? 
Around  Washington,  D.C.,  the  first  seems  to  be  crowded  onto  the  Cretaceous 
rock  formations,  leaving  H.  c.  cinerea  to  the  east  on  the  marl,  alum  bluff,  and 
limestones  of  the  Miocene  and  on  the  alluvial  sands. 

We  would  all  like  to  know  more  about  finding  the  delicate  little  Syrrhophus 
marnoctyi,  which  chooses  soft  soil  under  stone  or  log  shelter  for  habitat. 
Around  San  Marcos  and  Helotes,  Tex.,  are  Austin  chalk,  Eagle  Ford  shale, 
Edwards  or  Fredericksburg  limestone,  and  the  limestone  to  clay  of  the  Wash- 
ita  group  in  close  association.  In  Tom  Green  County,  where  this  frog  has 
been  found,  occur  similar  structures  with  the  possibility  of  San  Angelo  sand- 
stone  and  Blaine  gypsum  added.  In  Green  Gulch,  Chisos  Mountains,  we  again 
find  Eagle  Ford  shales,  Fredericksburg  limestone,  Chisos  sandstone  tuff, 
touches  of  Washita  group,  and  varying  amounts  of  Austin  chalk  in  close 
proximity.  Does  the  different  combination  help  vary  the  frogs9  structure  and 
pattern  a  bit,  giving  us  S.  gaigeae? 

What  does  the  robber  frog  (Eleutherodactylus  latrans)  like?  Its  distribu- 
tion is  surprising.  For  years  it  was  known  only  from  the  region  of  San  An* 
tonio,  Tex.,  and  from  Madera  Canyon  in  the  Santa  Rita  Mountains  in 


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jo  HANDBOOK  OF  FROGS  AND  TOADS 

Arizona,  a  separation  of  almost  1000  miles.  It  has  recently  been  found  in  New 
Mexico  near  Carlsbad,  a  limestone  and  gypsum  country,  and  is  reported  in 
Arizona  north  of  Roosevelt  Lake,  from  Parker  Canyon,  where  there  is  also 
limestone.  There  are  unusually  good  combinations  of  rock  conditions  around 
Helotes,  Hondo,  and  Uvalde,  Tex.,  to  make  desirable  homesteads  for  this  no- 
tional form.  The  breaks  of  the  Balcones  Fault  occur  at  the  front  of  the  lime- 
stones of  the  Edwards  plateau  allowing  water  to  come  to  the  surface,  and  there 
is  plentiful  underground  water.  The  several  kinds  of  limestone  conserve  the 
water,  and  gypsum  content  easily  washes  out,  leaving  caves,  cavities,  and  worn 
rock  ledges  readily  available  for  shelter. 

"Distribution  of  the  Flora  and  Fauna  of  Louisiana  in  Relation  to  Its  Geol- 
ogy and  Physiography"  (La.  Acad.  Set.,  8, 11-73  [Dec.,  1944])  shows  well  the 
interlocking  of  habitats,  and  Percy  Viosca,  Jr.,  explains  (p.  55)  how  two  forms, 
Acris  g.  gryllus  and  A.  g.  crepitans,  may  live  in  one  pond  separated  by  an  in- 
visible barrier,  the  one  at  the  inlet  where  acid  water  from  pinelands  is  present, 
the  other  at  the  outlet  where  the  alkaline  influence  of  the  tide  is  felt. 

Similar  intimate  studies  of  particular  spots  must  be  made  before  we  can 
understand  the  apparent  separation  into  broken  spots  and  the  apparent  meet- 
ing of  ranges  of  closely  related  forms. 

Habitat:  This  topic  usually  refers  to  nonbreeding  habitats,  but  at  times 
illusions  are  made  to  breeding  localities  as  well. 

Size:  The  phrasing  we  have  employed  is:  adults  3^-6%  inches  (males, 

$0-156  mm.;  females,  87-165  mm.).  These  are  the  measurements  of  the  large 

Colorado  River  toad,  Bufo  alvarius  Girard.  They  mean  that  breeding  adults 

•*  from  3%  to  6%  inches  in  length  of  body  from  tip  to  snout  to  rear  end 

1 1 .  fy  back  of  tbr     m*.  "i  ^  y/&  inches  or  80  mm.  is  the  smallest  size  at 

Ji'  maAv5  rhature,  and  6%  inches  or  165  mm.  is  the  largest  size  of  any 

.measured  female.  Almost  invariably  the  lower  measurement  in  inches  will 

be  that  of  a  male,  and  the  maximum  adult  measurement,  the  size  of  a  female. 

Jordan  and  Evermann  considered  the  two  killifishes,  Heterandria  formosa 
and  Lucania  ommata,  the  smallest  vertebrates  of  North  America.  We  have 
collected  many  of  these  tiny  fish,  but  confidently  pronounce  the  little  chorus 
frog,  Pseudacris  ocularis,  much  smaller.  Cuba,  we  understand,  has  a  still 
smaller  frog. 

Most  of  the  seven  families  have  extremes  in  size.  The  tree  frog  adults  vary 
from  %o  inch  (11.5  mm.)  in  the  little  chorus  frog  (P.  ocularis)  to  5%  inches 
(130  mm.)  in  the  Key  West  tree  frog  (Hyla  septentrionalis).  The  robber  frog 
adults  vary  from  %  inch  (15  mm.)  in  Camp's  frog  (Syrrhophus  campi)  to  3% 
inches  (90  mm.)  in  the  Texas  cliff  frog  (Eleutherodactylus  latrans).  The  adult 
toads  vary  from  %  inch  (19  mm.)  in  the  oak  toad  (Bufo  qucrcicus)  to  6% 
inches  (165  mm.)  in  the  Colorado  River  toad  (B.  alvarius).  The  adult  spade- 
foots  vary  from  i%  inches  (37.5  mm.)  in  Hammond's  spadefoot  (Scaphiopus 
A,  hammondii)  to  2%  inches  (72  mm.)  in  Holbrookes  spadefoot  (S.  holbroo\ii 


M  HANDBOOK  OF  PROGS  AND  TOADS 

holbroo\ii).  The  adult  frogs  vary  from  i%  inches  (36  mm.)  in  the  northern 
wood  frog  (Rana  s.  cantabrigensis)  to  8  inches  (200  mm.)  in  the  bullfrog 
(R.  catesbeiana) .  Two  groups  have  little  variation  in  their  extremes.  The 
narrow-mouthed  toads  vary  in  adults  from  %  inch  (20  mm.)  in  the  Texas 
narrow-mouthed  toad  (Microhyla  olivacea)  to  i%  inches  (41  mm.)  in  Taylor's 
toad  (Hypopachus  cuneus).  The  one  species  of  ribbed  toads  (Ascaphus  truei) 
varies  from  i%  to  2  inches  (28-51  mm.)  in  length. 

General  appearance:  In  most  cases  these  accounts  were  written  with  a  live 
specimen  or  specimens  in  hand.  Each  gives  the  form  of  body  (habitus)  of  the 
animal,  the  color  of  the  animal,  and  some  of  the  other  outstanding  characters. 
Often  the  animal  is  compared  to  a  closely  related  species  or  to  the  common 
type  of  the  group. 

Color:  A  consistent  effort  has  been  made  to  secure  a  color  description  from 
live  specimens  mainly  in  the  field  or  occasionally  in  the  laboratory.  Each  sex  is 
described.  It  must  be  remembered  each  description  is  of  a  particular  specimen. 
The  description  follows  the  Ridgway  code  (2d  ed.),  and  his  spelling.  Any  de- 
scription not  following  this  code  is  marked  "non-Ridgway." 

Structure:  This  section  is  meant  to  supplement  the  characters  given  under 
"General  appearance"  or  to  add  to  characters  used  in  the  keys.  Whereas  "Gen- 
eral appearance"  is  written  from  living  animals,  "Structure"  is  added  from 
examinations  of  preserved  specimens  and  from  published  descriptions.  Unless 
otherwise  stated,  all  measurements  in  descriptions  or  keys  are  relative  to  the 
body  length  (represented  by  L.).  "Structure"  is  written  in  the  abbreviated 
form  preferred  in  scientific  descriptions.  Very  frequently  the  original  descrip- 
tion appears  in  this  section. 

Voice:  Early  travelers  often  commented  on  the  frog  music  of  our  country. 
Witness  the  following: 

"There  be  also  store  of  frogs,  which  in  the  spring  time  will  chirp,  and  whistle 
like  birds;  there  be  also  toads,  that  will  creep  to  the  top  of  trees,  and  sit  there  croak- 
ing, to  the  wonderment  of  strangers!" 

"To  the  stranger  walking  for  the  first  time  in  these  woods  during  the  summer, 
this  appears  the  land  of  enchantment;  he  hears  a  thousand  noises,  without  being 
able  to  discern  from  whence  or  from  what  animal  they  proceed,  but  which  are,  in 
fact,  the  discordant  notes  of  five  different  species  of  frogs!" 

Previous  to  my  coming  to  this  country,  I  recollect  reading  the  foregoing  passages, 
the  first  in  a  history  of  New  England,  published  in  London,  in  the  year  1671;  and 
the  other  in  a  similar  production  of  a  later  date. 

Prepared  as  I  was  to  hear  something  extraordinary  from  these  animals,  I  confess 
the  first  frog  concert  I  heard  in  America  was  «o  much  beyond  anything  I  could  con- 
ceive of  the  powers  of  these  musicians,  that  I  was  truly  astonished.  This  perform- 
ance was  al  fresco,  and  took  place  on  the  night  of  the  i8th  instant  [April  18, 1794, 
Philadelphia],  in  a  large  swamp,  where  there  were  at  least  ten  thousand  perform- 
ers; and  I  really  believe  not  two  exactly  in  the  same  pitch,  if  the  octave  can  possibly 
admit  of  so  many  divisions  or  shades  of  semitones.  An  Hibernian  musician,  who,  like 


GENERAL  ACCOUNT  13 

myself,  was  present  for  the  first  time  at  this  concert  of  antimusic,  exclaimed,  "Be- 
gorrah,  but  they  stop  out  of  tune  to  a  nicety" 

I  have  been  since  informed  by  an  amateur,  who  resided  many  years  in  this  coun- 
try, and  made  this  species  of  music  his  peculiar  study  that  on  these  occasions  the 
treble  is  performed  by  the  tree-frogs,  the  smallest  and  most  beautiful  species;  .  .  . 
their  note  is  not  unlike  the  chirp  of  a  cricket:  the  next  in  size  are  our  counter  tenors; 
they  have  a  note  resembling  the  setting  of  a  saw.  A  still  larger  species  sing  tenor; 
and  the  under  part  is  supported  by  the  bull-frogs;  which  arc  as  large  as  a  man's  foot, 
and  bellow  out  the  bass  in  a  tone  as  loud  and  sonorous  as  that  of  the  animal  from 
which  they  take  their  name. — William  Priest,  Travels,  London,  1802,  pp.  48-50. 

Doubtless  all  males  have  voices,  yet  one  has  to  be  careful  in  such  records. 
One  of  the  phases  of  Pacific  Coast  frogs  and  toads  about  which  all  workers 
have  the  least  evidence  is  the  voice  and  vocal  sacs;  and  it  is  a  striking  fact  that 
Rana  boylii,  R.  aurora,  R.  pretiosa,  and  Bufo  boreas  have  no  vocal  sacs  appar- 
ent in  preserved  specimens  or  particularly  apparent  in  live  individuals.  Are 
they  like  some  eastern  forms,  namely,  R.  clamitans,  R.  catesbeiana,  and  JR.  syl- 
vatica? 

The  impression  that  Boulenger  gives  and  the  layman  has,  namely,  that  fe- 
males are  mute,  is  not  exactly  the  situation.  Let  anyone  pick  up  a  female  soli- 
tary spadefoot  (Scaphiopus  holbrootyi)  and  squeeze  it,  and  he  might  think 
he  had  a  male.  Those  who  know  the  species  would  recognize  the  difference 
between  the  male  and  the  female  voice.  Or  lay  this  same  female  on  her  back 
and  stroke  her  belly,  and  she  will  speak  vigorously,  possibly  not  so  strenuously 
as  a  male  but  nevertheless  the  voice  is  there.  All  spadefoot  females  can  and  do 
croak  some.  In  the  same  way  females  of  spadefoots,  frogs,  and  some  other 
groups  can  open  the  mouth  and  give  a  cry  or  scream  note  just  as  young  bull- 
frogs have  alarm  notes  as  they  go  skipping  to  cover.  Any  field  naturalist,  if  he 
has  heard  the  peculiar  cry  of  frogs  and  toads  when  in  the  grip  of  a  snake  or 
turtle,  very  well  knows  females  can  make  themselves  heard.  The  mercy  cry 
of  a  toad  or  frog  (like  jR.  catesbeiana,  R.  pipiens,  R.  p.  sphcnocephala,  R.  gry- 
lio,  or  some  Hylas)  can  be  and  is  as  likely  to  be  that  of  a  female  as  a  male. 
Miss  Dickerson  is  quite  right  in  saying  that  females  are  not  necessarily  voice- 
less. Doubtless  in  time  we  will  find  many  species  with  more  or  less  vocal 
females. 

Some  of  the  adjectives  employed  in  describing  American  frog  notes  are  bub- 
bling, weird,  plaintive,  hoarse,  woeful,  mournful,  complaining,  nasal,  inces- 
sant, musical,  pleasant,  whistling,  prolonged,  mellow,  tremulous,  squawking, 
shrill,  deafening,  ventriloquial,  peeping,  metallic,  resonant,  twittering,  loud, 
guttural,  snoring,  snorting,  gurgling,  clacking,  explosive,  grating,  and  sweet. 

Man  has  attempted  to  characterize  the  voices  of  frogs  since  Aristophanes, 
but  with  varying  success.  Voice  has  been  one  of  our  most  valuable  clues  on  life 
histories,  yet  we  must  say  it  is  hard  for  us  to  describe  a  call  as  others  do.  One's 
vehicle  of  description  may  be  figures  of  speech,  or  it  may  be  musical,  phonetic, 


14  HANDBOOK  OF  FROGS  AND  TOADS 

mechanical,  graphic,  or  biological.  Frog  voices  can  be  portrayed  by  tonal 
graphs.  They  are  also  presented  on  phonograph  records,  such  as  "Voices  ol 
the  Night"  (Comstock  Publishing  Company,  Inc.,  Ithaca,  New  York). 

Each  species  usually  has  its  distinctive  type  of  breeding  song,  yet  each  species 
may  have  several  variations.  Hyla  gratiosa  may  bark  in  the  high-  long-leaved 
pines,  but  "coatbet"  in  the  breeding  pools.  A  peeper  on  its  way  to  the  pond 
might  possibly  call  differently  from  one  with  the  chorus.  One  might  call  dif- 
ferently in  cold  or  warm  weather,  dry  or  rainy  spells.  The  breeding  song  is 
usually  typical,  but  one  needs  to  beware  of  believing  it  is  the  same  at  all  times, 
The  main  thing  is  to  learn  the  quality  of  the  song  in  order  to  be  ready  for  its 
variations.  Frogs  can  be  as  individualistic  in  this  regard  as  are  other  animals. 

It  is  a  rather  remarkable  fact  that  one  of  the  most  frequent  observations  of 
earlier  travelers  (100  or  150  years  ago)  in  this  country  was  on  the  frog  music 
of  North  America,  but  in  the  scientific  names  none  suggest  voice  except  in  the 
early  spring  peepers  (cricket  frogs  or  savanna  crickets,  Acris  gryllus,  A.  crepi- 
tans;  chorus  frogs  or  swamp  cricket  frogs,  Pseudacris)  and  in  three  of  the  larg- 
est frogs,  12.  clamitans,  R.  grylio,  and  R.  pipiens.  In  the  same  way  common 
names  suggesting  voice  are  not  frequent.  None  of  the  vociferous  spadefoots, 
toadsyand  narrow-mouthed  toads  have  common  names  suggesting  their 
voices. 
^Several  Hylas  and  others  have  come  to  be  recognized  as  rain  indicators. 

If  one  knows  the  frog  notes,  he  can  in  one  night  do  more  work  on  frog  dis- 
tribution than  he  might  otherwise  do  in  years.  In  one  favorable  night  of  auto 
riding  from  Lakehurst  through  southern  New  Jersey  we  located  many  colo- 
nies of  JR.  virgatipes  which  might  have  escaped  us  entirely  under  other  cir- 
cumstances. The  first  record  of  a  new  form  usually  comes  from  a  voice  record. 
We  never  would  have  found  our  first  Microhyla  near  Richmond,  Va.,  or 
H.  andersonii  and  JR.  virgatipes  on  an  overnight  camp  near  Everett  Pond, 
N.C.,  without  first  hearing  them.  This  was  our  method  near  San  Antonio  in 
1925,  namely,  to  go  along  the  roads  and  listen  until  midnight  or  later  and 
then  pursue  the  species  newly  heard  or  most  commonly  calling  that  night. 

The  choruses  of  the  Southeast  are  immense.  Sometimes  as  many  as  five  to 
eight  species  may  be  breeding  in  one  place.  At  a  distance  it  may  be  a  perfect 
din  but  to  wade  among  the  performers  one  encounters  a  deafening  concourse 
scarcely  imaginable.  Often  we  would  have  to  devise  methods  of  culling  out 
predominant  voices  to  catch  others  desired.  Sometimes  we  would  vary  the 
pressure  of  our  own  ear  openings  or  cup  our  ears  to  sift  one  particular  call  out 
of  the  chorus,  e.g.,  Hyla  fcmordis  would  drown  out  Bufo  quercicus. 

Vocal  sacs  (Plate  I) :  Doubtless  all  the  males  of  our  eastern  species  of  frogs 
do  croak  and  most  of  them  have  vocal  sacs.  Both  males  and  females  occasion- 
ally open  their  mouths  to  cry  when  in  distress  from  teasing,  alarm,  injury,  or 
capture,  but  all  croaking  males  of  the  United  States  and  Canada  keep  their 
mouths  closed  whether  in  air  or  under  water. 


Plate  L  Vocal  sacs.  i.  Southern  toad,  Bttfo  terrestris.  2.  Oak  toad,  Bufo 
quercicus.  3.  Northern  gopher  frog,  Rana  areolata  circulosa.  4.  Narrow-mouth 
toad,  Microhyla  carolinensis.  5.  Spadefoot  toad,  Bufo  compactilis.  6.  Spadefoot, 
Scaphiopus  holbroofyi  holbroofyi.  7.  Cricket  frog,  Acris  gryllus  gryllus.  8.  Little 
chorus  frog,  Pseudacris  ocularis.  9.  Pine  woods  tree  frog,  Hyla  jemoralis. 
10.  Barker,  Hyla  gratiosa.  n.  Gopher  frog,  Rana  capita.  12.  Southern  bullfrog, 
Rana  grylio.  13.  Sphagnum  frog,  Rana  virgatipes. 


16  HANDBOOK  OF  FROGS  AND  TOADS 

Two  of  our  students  have  made  a  study  of  vocal  sacs.  In  1931  Miss  Rachel  E. 
Field  made  a  detailed  portrayal  (unpublished  MS)  of  the  vocal  sacs  of  our 
eastern  forms,  and  in  1935  Dr.  C.  C.  Liu  examined  our  American  forms  and 
made  comparisons  with  world  forms.  Dr.  Lui  found  the  following: l 

Vocal  sacs  and  vocal  sac  openings  absent  (5) : 
Ascaphtfs  truei  Rana  boylii  muscosa 

Bufo  halophilus  Rana  b.  sierrae 

Rana  aurora  aurora 

Median  subgular  internal  sac  and  slitlike  openings  (17) : 
Scaphiopus  couchii  Bufo  hemiophrys 

Scaphiopus  hammondii  Bufo  marinus 

Scaphiopus  holbrookji  Bufo  punctatus 

Bufo  alvarius  Bufo  terrestris 

Bufo  americanus  Bufo  valliceps 

Bufo  boreas  Syrrhophus  campi 

Bufo  calif ornicus  Syrrhophus  marnoc1(ii 

Bufo  canorus  Hyla  femoralis 

Bufo  cognatus 

Median  subgular  external  sac  and  slitlike  sac  openings  (24) : 
Bufo  compactilis  Hyla  gratiosa 

Bufo  debilis  Hyla  regilla 

Bufo  fotvleri  Hyla  squirclla 

Bufo  quercicus  Hyla  versicolor  versicolor 

Bufo  woodhousii  Hyla  v.  chrysoscelis 

Acris  gryllus  Pseudacris  brachyphona 

Hyla  andersoni  Pseudacris  nigrita 

Hyla  arenicolor  Pseudacris  ocularis 

Hyla  avivoca  Pseudacris  ornata 

Hyla  baudinii  Pseudacris  triseriata 

flyla  cinerea  Microhyla  carolinensis 

'Hyla  crucifer  Microhyla  texensis 

Median  subgular  sac  internal  and  round  sac  openings  (i) : 

Hyla  septentrionalis 

Median  subgular  sac  external  and  round  vocal  openings  (o). 
Paired  subgular  sacs  internal  and  slitlike  openings  (o). 
Paired  subgular  sacs  external  and  round  openings  (o). 
Paired  subgular  sacs  internal  and  round  openings  (i) : 

Rana  palmipes 

Paired  lateral  sacs  internal  and  slitlike  openings  (2) : 
Rana  sylvatica  latircmis  Rana  septentrionalis 


1  The  names  are  those  used  by  Dr.  Liu  in  his  thesis. 


Plate  II.  Throat  differences  in  male  and  female,  i.  Cricket  frog,  Acris 
gryllus  crepitans,  male.  2.  Cricket  frog,  A.  g.  crepitans,  female.  3.  Narrow- 
mouth  toad,  Microhyla  carolinensis  t  male.  4.  Narrow-mouth  toad,  M.  carolinen- 
sis,  female.  5.  Eastern  chorus  frog,  Pseudacris  nigrita  feriarum,  female,  6. 
Eastern  chorus  frog,  P.  n.  jeriarum,  male.  7.  Spadefoot  toad,  Bufo  compactilis, 
male  (rear  throat  with  pleats  or  lappet).  8.  Southern  toad,  Bufo  terrestris, 
female.  9.  Southern  toad,  Z?.  terrestris,  male.  10.  Great  Plains  toad,  Bufo 
cognatus,  male  (rear  throat  dark),  u.  Barker,  Hyla  gratiosa,  male.  12.  Barker, 
H.  gratiosa,  female.  13.  River-swamp  frog,  Rana  hect(scherit  female.  14.  River- 
swamp  frog,  R.  hccl^cheri,  male. 


i8  HANDBOOK  OF  FROGS  AND  TOADS 

Paired  lateral  sacs  internal  and  round  openings  (9) : 
Rana  boylii  boylii  Rana  prctiosa 

Rana  catesbeiana  Rana  septentrionalis 

Rana  grylio  Rana  sphenocephala 

Rana  palustris  Rana  sylvatica 

Rana  pipiens 

Paired  lateral  sacs  external  and  round  openings  (5) : 
Rana  aesopus  Rana  draytonii 

Rana  areolata  Rana  montezumae 

Rana  clamitans 

These  two  were  careful  workers,  yet  we  wish  more  information  on  live 
males  to  answer  such  inquiries  as  the  following: 

1.  Why  should  Bufo  boreas  and  B.  b.  halophilus  be  in  different  categories? 
Or  Rana  aurora  and  R.  a.  draytonii?  Or  JR.  boylii  and  R.  b.  sierrae,  R.  b.  mus- 
cosa? 

2.  Why  is  Bufo  cognatus  out  of  the  B.  compactilis,  B.  debilis,  B.  quercicus 
class  ? 

3.  Why  is  Hyla  femoralis  unlike  our  other  Hylids  (except  H.  septentri- 
onalis) P 

4.  Why  is  R.  sylvatica  unlike  R.  s.  cantabrigensis  or  R.  s.  latiremis? 

Color  of  throat  (Plate  II) :  In  Ascaphus  truei,  Syrrhophus  campi,  S.  mar- 
nocf(ii,  Eleutherodactylus  ricordii,  E.  latrans,  Bufo  b.  borcas,  B.  b.  halophilus, 
R.  s.  cantabrigensis,  R.  aurora,  R.  boylii,  R.  pretiosa,  R.  sylvatica,  the  color  of 
the  throat  of  the  male  is  little  different  from  that  of  the  female.  These  are 
forms  with  no  prominent  vocal  sacs.  In  R.  palustris,  R.  pipiens,  R.  p.  spheno- 
cephala, R.  capito,  and  jR.  areolata,  forms  with  lateral  sacs,  only  slight  differ- 
ences occur  in  the  throat  color.  In  the  narrow-mouths  each  sex  may  have  a 
dark  throat.  Usually  the  male's  throat  is  darker,  but  it  is  often  difficult  to  use 
this  criterion.  Sometimes  in  Scaphiopus  holbroo1(ii  and  S.  couchii  the  differ- 
ence is  not  readily  recognizable.  In  S.  hammondii  enough  of  the  dark  slaty  or 
bluish  cast  appears  in  the  male's  throat  to  distinguish  it. 

The  pronounced  differences  in  throat  coloration  come  in  the  Hylidae  (ex- 
cept Pseudacris  ocularis),  Bufonidae,  and  Ranidae.  Only  seven  of  the  twenty- 
one  forms  of  Rana  have  this  marked  difference.  These  tend  to  have  yellows 
(sulphur  yellow,  maize  yellow,  sulphine  yellow,  oil  yellow,  lemon  yellow, 
olive  ocher,  aniline  yellow,  primuline  yellow,  or  barium  yellow)  in  the  males, 
and  in  the  females  lighter  washes  of  yellow  or  white;  e.g.,  the  green  frog  (R. 
clamitans)  male  has  a  barium  yellow  throat,  the  female  a  white  throat.  If  each 
has  the  same  yellow  the  male  may  have  dark  spots  or  some  other  equally  strik- 
ing character. 

The  whole  throat  of  some  species  of  toads  will  be  discolored  or  darker  than 
the  rest  of  the  venter,  or  only  the  lower  part  of  the  throat  may  be  thus  colored, 
as  in  the  sausage  group  (Bufo  cognatus,  B.  quercicus,  B,  compactilis). 


Plate  111.  Enlarged  tympana  in  male  Ranas  (seven  species),  i.  Green  frog, 
R.  clamitans,  male.  2.  Green  frog,  7?.  clamitans,  female.  3.  River-swamp  frog, 
R.  hcc^scheri,  male.  4.  River-swamp  frog,  R.  hcc\scheri,  female.  5.  Mink  frog, 
R.  septentrionalis,  male.  6.  Mink  frog,  R.  septentrionalis,  female.  7.  Sphagnum 
frog,  R.  virgatipes,  male.  8.  Sphagnum  frog,  R.  virgatipes,  female.  9.  Nevada 
frog.  R.  fisheri,  male.  10.  Nevada  frog,  R.  fisheri,  female,  n.  Southern  bullfrog, 
R.  grylto,  male.  12.  Southern  bullfrog,  R.  grylio,  female.  13.  Bullfrog,  R.  catesbci- 
anat  male.  14.  Bullfrog,  R.  catcsbeiana,  female. 


20  HANDBOOK  OF  FROGS  AND  TOADS 

In  the  Hylidae,  Pseudacris  (except  P.  ocularis),  Acris,  and  Hyla  have  the 
throat  dark  and  the  female  has  the  throat  white  or  like  the  rest  of  the  venter. 
In  the  group  to  which  Hyla  cinerea  cinerea,  H.  c.  evittata,  H.  gratiosa,  and 
H.  andersonii  belong,  the  males  have  the  whole  throat  or  most  of  the  throat 
dark  or  green,  but  the  females  have  green  on  either  side  of  the  throat  below 
and  ahead  of  the  angle  of  the  jaw.  In  the  two  subspecies  of  H.  cinerea  females 
the  color  varies  in  degree  of  extension  of  green  on  sides  of  throat.  In  H.  gra- 
tiosa females  the  green  extension  is  more  or  less  demarcated  on  the  throat  side 
with  a  prominent  white  border,  and  in  H.  andersonii  this  white  border  be- 
comes clearly  defined  in  females. 

In  general,  males,  being  more  active,  are  inclined  to  darker  colors  on  the 
dorsum  than  the  females.  In  a  mated  pair  this  is  often  quite  noticeable,  but 
even  in  those  not  mated  the  contrast  may  be  very  pronounced.  This  discrep- 
ancy is  marked  in  early  spring  breeders  but  also  obtains  for  later  breeder's.  So 
also  old  males  incline  more  to  self-color  (witness  Ran  a,  etc.)  than  the  females, 
which  may  be  more  spotted.  Or  if  a  pattern  be  common  to  each,  it  will  be 
bright  in  the  female  (e.g.,  Scaphiopus  holbrootyi  females  may  have  yel- 
lowish or  buffy  stripes  or  spots  on  back  and  the  males  may  be  almost  uni- 
form). 

Enlarged  tympana,  males  (Plate  III) :  The  seven  species  of  frogs  (Rana)  in 
which  the  throats  of  the  males  may  be  differently  colored  from  those  of  the 
females  have  even  more  marked  differences  in  the  tympana.  These  seven  spe- 
cies first  breed  at  about  41  mm.  for  R.  virgatipes,  44  mm.  for  R.  fisheri,  48  mm. 
for  jR.  scptentrionalis,  52  mm.  for  R.  clamitans,  82  mm.  for  R.  hecl(scheri  and 
R*  grylio,  and  85  mm.  for  R.  catesbeiana. 


BREEDING  SIZES 
Name  Males  Females 


mm.  mm* 
Ascaphidae 

Ascaphus  truei  29-43  28-51 

Scaphiopodidae 

Scaphiopus  h.  bombifrons  38-56  43~56 

Scaphiopus  h.  intermontanus  4°~59  42-63 

Scaphiopus  h.  hammondii  42-52  44~52-5 

Scaphiopus  h.  hurteri  43-73  44~82 

Scaphiopus  couchii  48-70  50-80 

Scaphiopus  holbroo1(ii  54*72 


GENERAL  ACCOUNT 


ax 


Name 


Males 


Females 


Bufonidae 

Bufo  quercicus 

19-30 

2O-32 

Bufo  debilis 

26-45 

31-52 

Bufo  insidior 

— 

— 

Bufo  punctatus 

40-68 

42-74 

Bufo  a.  copei 

40 

75 

Bufo  w.  jowleri 

41-74-5 

56-82 

Bufo  b.  nelsoni 

-  -1TT7**  *"•*" 

42-68 

46-89 

Bufo  terrestris 

42-82 

44-88 

Bufo  calif  ornicus 

44-58 

44-4,8 

Bufo  txsul 

44-59 

46-65 

Bufo  canorus 

46-64 

45-74 

Bufo  cognatus 

47-95 

60-99 

Bufo  compactilis 

52-78 

54-91 

Bufo  a.  americanus 

54-85 

56-110 

Bufo  vaUiceps 

53-98 

54-125 

Bufo  hemiophrys 

56-68 

56-80 

Bufo  w.  woodhousit 

56-99 

58-118 

Bufo  b.  boreas 

56-108 

60-110 

Bufo  b.  halophilus 

62-101 

60-116 

Bufo  alvarius 

80-156 

87-165 

Bufo  marinus 

-220 

Hylidae 

Acris  g.  gryllus 

15-29 

l6~33 

Acris  g.  crepitans 

17-30 

20-35 

Pseudacris  ocularis 

11.5-15.5 

12-17.5 

Pseudacris  n.  scptentrionalis 

1Q-32 

19-35 

Pseudacris  n.  clartyi 

20-29 

21-31 

Pseudacris  n.  nigrita 

21-28 

22-30 

Pseudacris  n.  feriarum 

21-30 

22-35 

Pseudacris  ».  triseriata 

21-32 

20-37-5 

Pseudacris  n.  verrucosa 

22 

3° 

Pseudacris  brimleyi 

24-28 

27-32-5 

Pseudacris  brachyphona 

24-32 

27-34 

Pseudacris  ornata 

25-35 

32-36 

Pseudacris  strecJ^eri 

25-41 

32-48 

JHyla  crucifer 

18-29 

20-33 

Hyla  squirella 

23-36 

23-37 

Name 

Males 

Females 

mm. 

mm. 

Hyla  femoralis 

24-37 

23-40 

Hyla  wrightorum 

24-38 

24-42 

Hyla  regilla 

25-48 

25-47 

Hyla  v.  phaeocrypta 

28P-33 

32?-36 

Hyla  avivoca 

28-39 

32-49 

Hyla  arenicolor 

29-53 

29-53 

Hyla  andersonii 

30-41 

3N7 

Hyla  v.  versicolor 

32-51 

33-6o 

Hyla  v.  chrysoscelis 

36-43 

35-49 

Hyla  c.  evittata 

36-47 

32-47 

Hyla  c.  cinerea 

37-59 

4I~63 

Hyla  baudinii 

44-71 

44-89 

Hyla  septentrionalis 

46-75 

52-5-95 

Hyla  gratiosa 

49-68 

50-68 

Leptodactylidae 

Eleuthcrodactylus  r.  planirostris 

15-25 

17~3° 

Syrrhophus  cam  pi 

J5-23 

J5-25-5 

Syrrhophus  gaigeae 

21 

28 

Syrrhophus  marnoclyi 

21-33 

22-39.5 

Leptodactyltis  labialis 

35? 

-49 

Eleutherodactylits  latrans 

48-74 

57-90 

Ranidae 

Rana  s.  cantabrigensis 

29-50 

34-56 

Rana  sylvatica 

34-60 

34-68  or  82 

Rana  b.  boylii 

39-^7 

40-75 

Rana  virgatipes 

41-63 

41-66 

Rana  fisheri 

44-64 

46-74 

Rana  b.  sierrac 

44-72 

48-84 

Rana  onca 

44-68 

51-84 

\Rana  a.  aurora 

44-63 

52-87 

Rana  p.  pretiosa 

44-75 

46-95 

Rana  p.  hiteiventris 

— 

— 

Rana  b.  muscosa 

45-€6 

47-81 

Rana  palustris 

46-64 

49-79 

Rana  septentrionalis 

46-71 

48-76 

$ 


Plate  IV.  Secondary  sexual  characters,  i.  Enlarged  thumb,  single  pad,  Rana  pipiens 
pipiens,  male.  2.  Horny  excrescences  on  thumb  and  first  finger,  Bujo  amcricanus  ameri- 
canus,  male.  3.  Enlarged  thumb,  double  pad,  Rana  boylii  muscosa,  male.  4.  Enlarged 
forearm,  Bufo  boreas  borcas,  male.  5.  Horny  excrescences  on  thumb  and  first  finger, 
Scaphiopus  holbroo\ii  holbrootyi,  male.  6.  Prepollex,  Hyla  septentrionalis.  7.  Plaited 
lateral  vocal  sac,  Rana  arcolata  circulosa,  male.  8.  Dimorphism  in  mating  pair,  male 
smaller,  darker,  female  larger,  lighter,  Hyla  crttctjcr  crucijcr.  9.  Horny  excrescences  on 
arm  and  fingers,  Bujo  alvanus,  male.  10.  Longitudinal  throat  plaits,  Pseudacris  nigrita 
]eriarum,  male.  n.  Double  throat  sac,  Hyla  baudinu,  male.  12.  Apron  over  concealed 
lower  throat  sac,  Bufo  cognatus,  male.  13.  Tail  and  thumb  pad,  Ascaphus  truei,  male.  14. 
Excrescences  on  thumb  pad  and  forearm,  A.  truei,  male.  15.  Anal  tube,  A.  truci,  female. 


HANDBOOK  OF  FROGS  AND  TOADS 


Name 

Males 

Females 

mm. 

mm. 

Rana  p.  sphenocephala 

49-78 

53-82 

Rana  cascadae 

50-58 

52-74 

Rana  p.  pipiens 

52-80 

52-102 

Rana  clamitans 

52-95 

58-100 

Rana  tarahumarae 

58 

-"3 

Rana  a  draytonii 

63-100 

58-136 

Rana  capita 

68-ioi 

77-108 

Rana  a.  areolata 

— 

— 

Rana  sevosa 

-84 

-92.5 

Rana  grylio 

82-152 

85-161 

Rana  hecT(scheri 

82-131 

102-131 

Rana  catesbeiana 

85-180 

89-200 

Microhyla  olivacea 
Microhyla  carolinensis 
Microhyla  areolata 
Hypopachus  cuneus 


Brevicipitidac 


20-28 
20-30 
24-28 
28-37-5 


19-29.5 
22-36 
23-30 
36- 


SUMMARY  OF  SECONDARY  SEXUAL  CHARACTERS 

(Plate  IV) 

A.  Thumb  enlarged  in  male  (Ranidae  and  H.  septentrionalis) 
B.  Enlarged  tympana  (Rana  clamitans  group) 

C.  Males  have  external  vocal  sacs;  males  41-71  mm.,  females  41-76 
mm.  (Rana  virgatipes  group:  R.  fisheri,  R.  septentrionalis t  R. 
virgatipes) 

CC.  Males  without  external  vocal  sacs;  males  first  breeding  at  82-85 
mm.,  females  82-89  mm.,  except  JR.  clamitans  males  52-95  mm., 
females  58-100  mm.  (Rana  catesbeiana  group :  R.  catesbeiana, 
R.  clamitans,  R.  grylio,  R.  hecJ(scheri) 
BB.  No  enlarged  tympana 

C.  Base  of  thumb  swollen  but  no  prethumb  expansion  (prepollex) 
D.  No  external  vocal  sacs 

E.  Thumb  pad  in  two  parts  with  oblique  groove  (Rana 
boylii,  3  subsp.,  and  R.  aurora  group  somewhat,  3 
forms) 


GENERAL  ACCOUNT  25 

EE.  Thumb  pad  or  enlargement  not  in  two  parts;  web 
of  hind  foot  of  male  more  or  less  convex  (Rana  syl- 
vatica group:  R.  s.  sylvatica,  R.  s.  cantabrigensis, 
R.  p.  pretiosa) 
DD.  External  lateral  vocal  sacs;  webbing  margin  of  hind  foot 

not  modified  in  male 
CC.  Base  of  thumb  enlarged  with  a  striking  prethumb  expansion 

or  prepollex  (Hyla  septentrionalis) 
AA.  Thumbs  not  enlarged  nor  ball-like  nor  padlike  at  base 

B.  Black  or  dark  excrescences  on  first,  second,  and  third  fingers 

C.  "Tail"  in  male;  excrescences  on  breast  and  inner  edge  of  fore- 
arm; fore  limb  and  hind  limb  enlarged,  and  hind  foot  also  in 
male  (Ascaphus  truet) 
CC.  No  "tail"  nor  excrescences  on  breast  or  forearm 

D.  No  median  vocal  sac  or  darkened  throat  in  males,  with 
heavy  forearms  (Bufo  boreas  group;  B.  b.  boreas,  Z?.  b. 
halophilus) 
DD.  Median  vocal  sac  present  in  male 

E.  Vocal  sac  in  lower  throat  (in  alcohol,  apron  or  bib- 
like),  darkened  in  male  (Bufo  cognatus  or  sausage 
group :  B.  cognatus,  B.  compactilis,  B.  quercicus) 
EE.  Vocal  sac  not  principally  from  lower  throat 

F.  Throat  slightly  discolored  and  finger  ex- 
crescences not  very  prominent  (Bufo  punc- 
tatus  group:  B.  punctatus,  B.  debilis) 
FF.  Excrescences  normal  and  throat  discolored 
(Bufo  americanus  group:  B.  a.  americanus, 
B.  w.  fowleri,  B.  terrestris,  B.  w.  woodhousii, 
B.  hemiophrys,  B.  valliceps) 
FFF.  Arm  broader  in  male;  excrescences  cover 
whole  of  upper  fingers  and  first  finger  ex- 
crescence extends  onto  wrist  (Bufo  alvarius) 
FFFF.  Excrescences  on  top  of  fingers,  not  inner 
edge    primarily;    throats    little    discolored 
(Scaphiopus  group) 
BB.  No  black  or  dark  excrescences  on  fingers 

C.  Throat  of  males  with  median  sac — revealed  in  alcohol  by 
wrinkles,  folds,  plaits,  or  darker  color  (Hylidae,  Leptodacty- 
lidae) 
D.  Throat  not  discolored  in  male  or  particularly  unlike  that 

of  female  (Leptodactylidae,  5  species) 

DD.  Throat  discolored  in  male  or  particularly  unlike  that 
of  female  (Hylidae:  Acris,  Pseudacris,  Hyla) 


26  HANDBOOK  OF  FROGS  AND  TOADS 

E.  Distinctly    discolored     (Acris,    Pseudacris,    some 

Hylas) 
EE.  Not  so  distinctly  colored  in  male  (Hylas) 

F.  Females  with  green  on  throat  (Hyla  cinerea 
group) 
G.  Green  edged  inside  by  white  line  (Hyla 

gratiosa,  H.  andersonif) 
GG.  Green  not  so  edged  (Hyla  cinerea  cinerea, 

H.  c.  evittata) 

FF.  Females  with  throat  like  rest  of  venter  (Hyla 
avivoca,  H.  arenicolor,  H.  crucifer,  H.  femo- 
ralis,  H.  squirella,  H.  versicolor) 
DDD.  Throat  discolored  in  male  and  female,  less  so  in  latter 

(Microhyla) 
CC.  Males  with  paired  vocal  sacs  below  angles  of  jaw  (H.  baudi- 

nif) 

Ovulation  (Plates  V-IX) :  The  vast  bulk  of  frog  species  are  in  the  South 
and  Southwest,  the  warmer  climes.  In  the  North  activity  begins  with  the  going 
out  of  ice  or  the  first  warning  of  spring  and  moisture.  Most  of  our  texts  have 
been  written  for  the  northeastern  states  where  few  species  occur,  and  so  many 
writers  have  concluded  that  all  frogs  breed  in  early  spring  because  Rana  syl- 
vatica,  R.  pipiens,  and  Hyla  crucifer  do.  In  the  same  way,  many  thought  all 
salamanders  breed  in  early  spring  because  Ambystoma  maculatttm,  A.  jeffer- 
sonianum,  and  A.  tigrinum  do. 

It  is  hard  to  classify  species  on  one  set  of  observations  in  one  locality  or  re- 
gion. A  species  may  be  a  short,  early  breeder  in  the  northeastern  states,  e.g., 
Rana  pipiens,  yet  lay  eggs  in  July  in  Devil's  River,  Tex.  A  bullfrog  may  lay 
normally  in  June  or  July  but,  under  peculiar  circumstances,  lay  in  February 
in  San  Antonio,  Tex.  A  R.  sphenocephala  might  breed  early  in  February  or 
almost  any  month  in  the  South,  and  we  might  think  it  would  be  restricted  in 
the  North;  yet  we  find  it  breeding  in  Wabash  Valley,  111.,  in  September.  Even 
in  the  North  Bufo  americanus,  B.  jowleri,  Scaphiopits  holbrootyi,  and  R. 
clamitans  stretch  the  ovulation  periods — a  condition  more  normal  in  the  South 
and  Southwest. 

The  spadefoots,  toads,  small  tree  frogs  such  as  Acris  and  Pseudacris,  some 
larger  tree  frogs  (Hyla  squirella,  H.  jemoralis,  H.  crucifer,  H.  regilla),  and  a 
few  Ranas  spawn  in  transient  pools,  impermanent  situations,'  roadside  ditches, 
and  temporary  floodlands,  and  often  the  loss  must  be  immense.  In  the  North 
Rana  sylvatica,  R.  s.  cantabrigensis,  and  R.  pipiens  masses  may  be  caught  by 
the  surface  ice  of  spring  freezes.  Next  to  the  losses  in  toads  and  spadefoots, 
we  believe  the  species  with  small  surface  films  suffer  the  most  from  complete 
drying  up  of  pools,  thus  being  hung  up  by  evaporation  above  a  pond's 
level. 


GENERAL  ACCOUNT 


Plate  V.  Eggs:  spadefoots,  narrow-mouth  toads  (Scaphiopus,  Microhyla). 
1,8.  Hammond's  spadefoot,  5.  hammondii  hammondti.  2,4.  Spadcfoot,  S. 
holbroot(it  holbrooktt.  3.  Couch's  spadefoot,  S.  couchii.  5.  Texas  narrow-mouth 
toad,  M.  olivacea.  6,7.  Narrow-mouth  toad,  M.  carolinen$i$. 


28 


HANDBOOK  OF  FROGS  AND  TOADS 


Plate  VI.  Eggs:  toads  (Bujo).  1,2,5.  Oak  toad,  B.  qncrcicus.  3,8.  Nebulous 
toad,  B.  valliceps.  4,10.  Spotted  toad,  B.  punctatus.  6.  Southern  toad,  B.  ter- 
restris.  7.  American  toad,  B,  americanus  americanus.  9,  Spadefoot  toad,  B. 
compactilis. 


GENERAL  ACCOUNT 


Plate  VII.  Eggs:  cricket  frog,  chorus  frog,  tree  toad  (Acris,  Pscudacris,  Hyla). 
i.  Little  chorus  frog,  P.  ocularis.  2,5.  Eastern  chorus  frog,  P.  nigrita  feriarum. 
3.  Cricket  frog,  A.  gryllus  gtyllus.  4.  Clarke's  chorus  frog,  P.  n.  clarfyi.  6.  Striped 
tree  frog,  P.  n.  t riser iata.  7.  Peeper,  H.  crucijer  cructfer.  8.  Squirrel  tree  frog, 
H.  squire/la.  9.  Green  tree  frog,  H.  cincrca.  10,13.  P^nc  woods  tree  frog,  H. 
femoralis.  ir.  Common  tree  toad,  f/.  versicolor  versicolor.  12,14.  Barker,  H. 
gratiosa. 


HANDBOOK  OF  FROGS  AND  TOADS 


Plate  VIII.  Eggs:  frogs  (Ratta).  i.  Sphagnum  frog,  R.  virgatipcs.  2.  Gopher 
frog,  R.  capita.  3.  Southern  bullfrog,  R.  grylio.  4.  Pickerel  frog,  R.  palustris.  5. 
California  yellow-legged  frog,  R.  boyln  boyhi.  6.  Nevada  spotted  frog,  R.  pretiosa 
luteiventns.  7.  Bullfrog,  R.  catesbeiana.  8.  Wood  frog,  R.  sylvatica  sylvatica.  9. 
Green  frog,  R.  cl ami  tans. 


GENERAL  ACCOUNT  31 

Egg-laying  process:  The  egg  forms  may  be  summarized  as  follows: 

(1)  Single  eggs  each  singly  ovulated  after  movement  of  pair,  e.g.,  Hyla 
crucifer. 

(2)  Single  eggs,  several  strewn  at  one  time  followed  by  movement,  e.g., 
several  Hylas. 

(3)  Single  eggs  laid  as  in  (2)  or  small  egg  packets  at  surface,  then  move- 
ment by  pair,  e.g.,  Hyla  cmerea. 

(4)  Small  packets  of  eggs  ovulated  at  one  time  at  the  surface,  movement  by 
pair,  e.g.,  Hyla  versicolor. 

(5)  Small  packets  of  eggs  ovulated  at  one  time,  no  movement  of  pair  and 
a  resultant  large  film  at  the  surface,  e.g.,  Rana  catesbetana. 

(6)  Eggs  laid  in  two  rosarylike  strings,  e.g.,  Ascaphus  truei. 

(7)  Files  (three-  or  two-  or  one-rowed),  movement  from  place  to  place  by 
pair,  e.g.,  Bufos. 

(8)  Bands,  later  swollen  to  cylinders,  regular  or  irregular,  like  two  films 
coalesced  or  two  oviducal  emissions  merged;  movement  from  bottom  or  base 
of  stem  to  tip,  e.g.,  Scaphioptts. 

(9)  Balls,  lumps,  spheres,  or  plinths,  merged  oviducal  emissions;  mainte- 
nance of  one  position  by  pair  during  entire  process  or  for  many  emissions,  e.g., 
meadow  frogs,  gopher  frogs,  wood  frogs,  Pacific  Coast  frogs;  Rana  virgattpes 
and  R.  septentrionahs. 

The  series  is  not  so  simple  as  it  is  given  above  because  of  movement  or  non- 
movement,  surface  or  submerged  laying,  size  of  frog,  egg  complement,  and 
season. 

Eggs:  The  species  of  the  United  States  and  Canada  whose  eggs  are  not  re- 
corded are  Scaphtoptis  holbrootyi  albus,  Bttfo  caltformcus,  B.  exsul,  B.  hemi- 
ophrys,  Hyla  avivoca,  H.  bauchnn,  Syrrhophus  campi>  S.  gatgeae,  S.  rnarnocfyi, 
Rana  boyltt  mitscosa,  R.  hecfyscheri,  R.  fishert,  R.  onca,  Mtcrohyla  areolata, 
M.  mazatlanensis — 15  in  all.  Breeding  notes  have  been  made  but  egg  descrip- 
tions are  lacking  for  Bttfo  boreas  boreas,  B.  debilis,  Pseudacris  ornata,  P.  nignta 
nigrita,  P.  n.  septentnonahs,  Hyla  ctnerea  evittata,  H.  wrightorum,  H.  versi- 
color chrysoscelis,  Eletitherodactylus  latrans,  Rana  areolata  areolata,  R.  syl- 
vatica  cantabrigensis,  R.  cascadae — 12  in  all.  Of  our  99  or  more  forms  we  thus 
have  27  species  whose  eggs  and  ovulation  habits  need  more  attention.  In  this 
country  we  know  the  eggs  of  about  70  species  minutely.  Of  these  almost  50 
species  have  been  seen  in  the  field. 

In  the  synopsis  of  egg  characters  we  have  drawn  in  part  on  the  work  of  Dr. 
Tracy  I.  Storer  for  Rana  aurora  draytontt,  R.  boyht  boyltit  Bufo  boreas  ha- 
lophilus,  Hyla  arenicolort  and  H.  regilla.  We  have  supplemented  our  observa- 
tions on  Scaphioptts  couchii  and  S.  hammondii  with  Dr.  Ortenburger's 
records.  For  Ascaphus  and  to  some  extent  Syrrhophus  marnocl(ii  notes,  we 
have  used  Mrs.  Helen  Thompson  Gaige's  collections  and  observations.  E.  R. 
Dunn's  and  R.  F.  Deckert's  studies  on  Eleutherodactylus  ricordii  and  Noble 
and  Noble's  work  on  Hyla  andersonii  have  been  incorporated  in  the  synopsis. 


32  HANDBOOK  OF  FROGS  AND  TOADS 

Relatively,  the  egg  vitelli  of  the  robber  frogs  (Syrrhophus  campi  and  S. 
marnoctyi,  3-4  mm.  in  diameter  for  yolk),  in  which  the  whole  development 
occurs  in  the  eggs,  are  the  largest,  with  Ascaphus  next.  Actually,  the  largest 
vitelli  are  those  of  the  bell  toad,  Ascaphus  truei,  in  which  yolks  of  5  mm.  occur. 
The  smallest  are  those  of  the  tiny  swamp  cricket  frog,  Pseudacris  ocularis,  with 
vitelli  0.6-0.8  mm.  in  diameter. 

The  size  of  the  adult  or  parent  does  not  determine  the  size  of  the  egg  or, 
more  strictly,  its  vitellus.  The  bullfrog,  the  largest,  has  vitelli  1.2-1.7  mm->  little 
larger  than  those  of  the  narrow-mouthed  toad,  Microhyla  carolinensis  (1.0-1.2 
mm.)  or  of  the  tree  toad,  Hyla  arenicolor,  and  smaller  than  those  of  Ricord's 
frog  (Eleutherodactyltts  ricordii  plantrostris,  2  mm.).  The  bullfrog  eggs  are 
only  0.6-0.9  mm-  Digger  than  those  of  our  smallest  species,  Pseudacris  ocularis. 
Within  one  group  wide  variation  takes  place.  Why  has  the  canyon  tree  toad 
(Hyla  arenicolor)  vitelli  1.8-2.4  mm.,  almost  twice  those  of  its  relative,  H.  fe- 
moralis,  0.8-1.2  mm.?  Or  why  should  one  of  the  smaller  frogs,  the  wood  frog 
Rana  sylvatica,  have  eggs  1%  times  larger  than  the  largest,  the  bullfrog?  In 
the  Northeast  we  concluded  (in  1914)  that  the  bullfrog  had  the  smallest 
vitellus  in  proportion  to  its  adult  size  and  that  Pseudacris  n.  triseriatus  and 
Rana  sylvatica  had  relatively  the  largest  vitelli.  In  the  Okefinokee  region, 
Geo.  (in  1929),  we  concluded  that  Bttfo  quercicus,  Pseudacris  ocularis,  P.  n. 
nigrita,  Acris  grylltts,  and  Microhyla  carolinensis  were  largest  in  vitelli  rela- 
tively, and  Rana  grylio  and  1?.  catesbeiana  were  the  smallest. 

The  season  of  breeding  for  species  in  the  North  has  its  beginning  and  end 
clearly  imrked.  Each  species  needs  four  or  five  weeks,  except  Btifo  a.  amcri- 
cantis  and  Rana  clamitans.  The  exceptions  may  require  two  or  three  months 
for  ovulation.  In  the  southeastern  states,  when  once  a  species  has  begun,  its 
season  of  breeding  may  extend  throughout  the  summer  or  even  into  the  early 
fall,  depending  upon  the  high  crests  of  precipitation.  These  species,  although 
of  a  swampy  region,  wait  for  the  rains,  and  in  this  reliance  on  precipitation 
they  suggest  our  desert  species  of  Texas  and  Arizona.  Those  that  do  not  begin 
until  June  have  at  least  eight  to  ten  weeks  of  ovulation.  This  minimum  period 
for  a  species  of  the  South  is  the  maximum  period  for  a  northern  form.  Species 
such  as  R.  p.  sphenocephala,  B.  terrestris,  and  Acris  gryttus,  which  begin  early 
in  the  season,  breed  during  25-30  weeks  of  the  year,  if  not  longer,  or  from  Feb- 
ruary to  October  or  November. 

When  one  comes  into  a  town  in  the  Big  Bend  country  of  Texas  or  in  the 
desert  region  of  Arizona  during  a  rain,  after  a  dry  period  of  six,  nine,  fif- 
teen, or  more  months,  one  wonders  how  it  is  that  spadefoots  and  toads  are 
ready  that  instant  for  their  congress  of  mating  and  ovulation.  We  can  only 
explain  it  by  supposing  that  the  ovulation  period  of  these  species  extends  over 
the  whole  spring  and  summer  and  that  all  individuals  are  not  ready  at  one 
particular  moment,  those  appearing  being  only  a  small  part  of  the  population. 
Perhaps  the  frogs  aestivate  or  hibernate  with  life  processes  slow,  and  the  eggs 


GENERAL  ACCOUNT  33 

in  this  condition  may  linger  almost  ready  for  ovulation  for  a  long  period. 

In  the  mountainous  regions  of  the  Far  West  the  season  of  ovulation  is  dic- 
tated by  the  short  term  of  four  or  more  months.  Forms  like  Rana  b.  sierrae  and 
Bujo  canorus  have  a  very  short  season  for  ovulation  and  tadpole  development. 
If  B.  canorus  did  not  come  from  a  group  of  short  larval  development — small 
tadpoles — we  would  not  be  surprised  at  its  wintering  over  as  larvae,  as  must 
some  1?.  b.  sierrae.  In  the  same  way  one  wonders  if  some  larvae  of  R.  s.  canta- 
brigensis  in  the  North  may  not  be  occasionally  forced  into  this  same  condi- 
tion. In  the  North  ice  goes  out  so  late  that  ovulation  and  development  periods 
are  considerably  restricted. 

The  number  ot  eggs  in  a  complement  may  vary  from  6  in  Syrrhophus  campi 
and  S.  marnocfyi,  or  35-51  in  the  bell  toad,  or  100  in  the  smallest  species, 
Pseudacrts  ocularis,  to  20,000  in  Rana  catesbciana,  the  largest  form.  The  range 
in  tree  frogs  (Hylulae)  is  from  100  (P.  oculans)  to  2084  (Hyla  gratiosa) ;  in 
the  toads  (Bufonidae)  from  610  (Bufo  quercictis)  to  16,500  (B.  b.  halophilus) ; 
in  the  frogs  (Ranidae)  from  349  (R.  virgatipes)  to  20,000  (R.  catesbeiand) . 
The  complements  of  the  narrow-mouthed  frog  (Microhyla  carolinensis)  and 
the  spadefoot  (Scaphioptts  holbroofyii)  are,  respectively,  869  and  2332. 

The  eggs  of  eight  species,  Hyla  cinerea,  H.  femoralis,  H.  verstcolor,  Rana 
catesbeiana,  R.  clamitans,  R.  grylio,  Microhyla  carolinensis,  and  M.  olivacea, 
float  on  the  surface  of  the  water;  the  eggs  of  30  or  more  species  (Acrist  Pseuda- 
crisf  Ascaphus,  Scaphiopus,  Bttfo,  14  species  of  Rana)  are  submerged.  In 
northern  or  southeastern  states  no  form  with  buoyant  eggs  lays  before  May  10, 
yet  in  Texas  Microhyla  ohvacea  may  breed  in  March  or  Rana  catesbeianat 
rarely,  in  mid-February.  The  19  or  20  normally  early  breeders  have  submerged 
eggs.  These  eggs  usually  have  firm  jelly  envelopes,  except  those  of  Pseudacns 
and  Scaphiopus ,  which  have  a  consistency  intermediate  between  the  firm  jel- 
lies of  early  breeders  and  the  loose  surface  films  of  late  breeders.  One  form, 
M.  carolinensis,  although  it  lays  at  the  surface,  has  the  most  beautifully  dis- 
tinct, firm  eggs  of  all  the  species  considered,  yet  its  Texan  relative,  M.  olivacea, 
has  indistinct  loose-jellied  eggs. 

We  had  eggs  laid  in  camp  and  in  the  laboratory  by  mated  pairs  caught  in 
the  field.  Later  the  eggs  in  the  field  were  determined  by  these  original  checks. 
Whenever  possible  the  process  of  egg  laying  was  observed  in  the  field.  Two 
species,  one  of  the  North  and  one  of  the  South,  were  identified  by  the  positive 
elimination  of  all  the  other  resident  forms. 

The  measurements  and  color  descriptions  are  based  on  the  fresh  eggs  of 
average  adults,  not  of  very  young  females.  Later  these  forms  were  checked 
with  preserved  material.  For  the  eggs  with  loose  outer  envelope  the  outer  mar- 
gin is  indicated  by  dots  (Plate  IX).  In  one  species  the  vitelline  membrane  is 
far  separated  from  the  vitellus,  and  the  space  is  indicated  by  cross  hatching. 
A  summary  of  the  egg  characters  of  each  species  follows  in  the  accompanying 
synopsis  (pp.  36-46). 


21 


I6o 


22 

T- 

>» 
23 


I 


/ 


I6b 


9o 


12 


V    / 


•  7o 


24 


25 


I7b 


9b 


7o 


26 


© 


27o 


27b 


7b 


10 


15 


28 


Plate  IX.  Diagrams  of  eggs  ty^2V>).  From  R.  L.  Livezey  and  A.  H.  Wright,  Amencan 
Midland  Naturalist,  37,  213-216  (1947).  i.  Eleutherodactyhts  ncordii  plamrostrts.  2. 
Ascaphus  truei.  3.  Scaphiopus  couchn.  4.  Scaphiopus  hatnmondn  hammondn,  the  only 
species  of  Scaphiopus  with  two  envelopes.  5.  Scaphtopus  holbroofyi  holbroofyi.  6.  Scaphi- 
opus holbtoo\n  hutteni.  7a,b.  B«/o  alvarms.  Note  odd  wedge-shaped  vitelh  in  b,  a  com- 
mon condition  in  this  species.  8.  Bufo  amcncanus  americanus.  9a,b.  Bufo  boreas  halo- 
philus.  10.  Bufo  calijormcus.  n.  Bujo  cognatus.  12.  Bufo  compactilis.  13.  Bufo  pu  net  at  us. 
The  only  Bufo  that  deposits  eggs  singly.  14.  Bufo  quetctcus.  15.  Bufo  tenestrts.  i6a,b. 
Bujo  valhceps.  I7«i,b.  Bujo  woodhousn  jowlert.  18.  //y/«  andersomt.  19.  //y/«  atemcolor. 
20.  //y/tf  cmerca  cineiea.  21.  f/y/^  cmcijcr.  22.  //y/«  jemorahs.  23.  Hy/0  gratiosa.  Vitelhne 
capsule  far  removed  from  vitellus.*  24.  f/y/j  regtlla.  25.  //y/a  squtrella.  26.  Hyla  versicolor 
versicolor.  27a,b.  -4rrw  gryllus.  Rarely  laid  as  in  b.  28.  Pseudacris  brachyphona. 


29 


31 


32o 


\ 


32b 


ZZc 


33         34 


35 


36 


\ 


45 


57  58 


29.  Pseudacns  mgnta  clartyi  30.  Pseudacns  n  jenarum.  31.  Pseudacns  n.  septcntnonalis. 
32a,b,c.  Pseudacns  n.  tnsenata.  Figure  a  is  probably  the  true  appearance  of  the  egg;  c  is 
after  Smith's  (1934)  description;  b  is  a  composite  of  a  and  c.  33.  Pseudacns  n.  venucosa. 
34.  Pseudacns  oculans.  35.  Pseudacns  ornata.  36.  Pseudacns  strec\en.  37.  Rana  aurora 
aurora.  38.  Rana  areolata  ctrculosa.  39.  Rana  aurora  dray  to  nit.  40.  Rana  boyln  boy  In. 
41.  Rana  b.  stctrae.  42.  Rana  cascadae.  43.  Rana  clamitans.  44.  Rana  capito.  45.  Rana 
catesbetana.  46.  Rana  gryho.  47.  Rana  palustns.  48  Rana  pipiens  ptpiens  49.  Rana  p. 
sphenocephala.  50.  Rana  pretiosa  pretiosa.  51.  Rana  p.  luteiventns.  52.  Rana  septen- 
trionalts.  53.  Rana  sylvattca  sylvattca.  54.  Rana  s.  cantabngensts.  55.  Rana  vtrgattpes. 
56.  Microhyla  carolmensis  carohnensts.  57.  Mtcrohyla  c.  olwacca.  58.  Hypopachus  cuneus. 
*  No  vitelhne  capsules  are  shown,  except  in  this  figure,  Hyla  grattosa. 


HANDBOOK  OF  FROGS  AND  TOADS 


SYNOPSIS  OF  EGGS  OF  UNITED  STATES  FROGS2 

A.  Eggs  unpigmcnted. 

a.  Laid  in  water  Rosary  like  string  in  circular  mass,  yolk  5  mm.  in 
diameter,  capsule  5  8  mm  A  tadpole  stage.  Egg  complement  35-49. 
Season  May  to  Sept.  11.  Ascaphus  truti 

aa.  Laid  on  land  or  in  moist  situations.  (Whole  development  within 
egg,  no  larval  stage  )  (No  data  for  Eleuthtrodactylus  latrans.) 
b.  Egg  complement  small,  6-10. 

c.  Complement  6  or  7.  Yolk  3-3.5  mm.,  av  3  o.  Season  April  to 
May.  Ovarian  evidence.  Syrrbopbus  campi 

cc.  Complement  fr-io,  yolk  about  4  mm.  Season  April  to  June  or 
July.  Ovarian  evidence.  Syrrhophus  marnockii 

bb.  Egg  complement  large,  11-15.  (R.  F.  Deckert  and  Dunn.)  Yolk 
i  mm.  in  diameter,  outer  envelope  eventually  4  mm.  Season 
April  to  August.  Elcuthcrodactylus  ricordii  planmstris 

PlateLXXVIII-5,7 

A  A.  Eggs  pigmented  (upper  dark  -pole  and  lower  light  pole).  Laid  in  water. 

B.  Eggs  deposited  singly 
a.  Envelopes  two. 

b  Outer  envelope,  diameter  1.4-1.0  mm.,  inner  envelope,  diameter 
1.  1-1.6  mm.,  vitellus,  diameter  o  8-1  o  mm.,  eggs  brown  above 
and  cream  below.  Egg  complement  941.  Season  June  10  to  Aug.  n. 

Hyla  squinlla 
Plates  VII-8,  IX-i5 

bb.  Outer  envelope,  diameter  3.5-4  o  mm.;  inner  envelope,  diameter 
i.gr  i.o  mm.,  vitellus,  i  1-1.4,  vitellus  dark  brown  and  creamy 
white,  strewn  among  sphagnum.  Egg  complement  800-1000. 
Season  May  i  to  July  10.  Hyla  andersonii 

aa.  Envelope  single. 

b.  Envelope  1.3  mm.  or  more. 

c.  Envelope  6.7  mm.  more  or  less,  loose,  sticky,  eggs  single, 
tandemhke,  often  Bufo  fashion  or  lattice  work  or  irregular. 
Vitelh  1.3  mm.,  black  above  and  white  or  gray  below.  Season 
April  to  July.  Scaphtopus  b.  burterit 


R    L.  Livczcy  and  A  H  Wright,  "Synoptic  Key,"  The  American  Midland 
Natural*  \t>  37,  179-111  (1947). 


GENERAL  ACCOUNT  37 

cc.  Envelope  1.3-5.0  mm.;  not  Bufo  fashion,  frequently  single  and 
irregularly  arranged, 
d.  Vitellus  0.9-1  8  mm. 

e.  Vitelline  membrane  far  from  vitellus,  appearing  as  inner 
envelope  i. 6-1.0  mm.,  outer  envelope  loose,  glutinous, 
indefinite  in  outline,  13-5  mm.,  vitellus  1.0-1.8  mm. 
Egg  complement  1084.  Season  March  3  to  Aug.  n. 

Hyla  grattosa 
Plates  VII-n,i4,  IX-i3;  LXIX-6 

ee.  No  inner  envelope  or  appearance  of  one,  envelope  firm, 
indefinite  in  outline,  1.3-3.6  mm. 

f.  Vitellus  0.9-1  o  mm.,  upper  pole  deep  brown,  buffy 
olive  lower  pole,  rarely  the  appearance  of  an  inner 
envelope.  Egg  complement  141.  Season  April  15  or 
earlier  to  Sept.  i.  Sometimes  in  masses. 

Acns  £.  gryllus 

Plates  VII -3,  IX  i7a,b. 

Acns  &.  crtpitam 

ff.  Vitellus  i  o  1.3  mm.,  upper  pole  black,  lower  pole 
white,  envelope  3  i  3.6  mm.  Egg  complement  1000 
to  3000.  Season  April  to  September.  Bufo  punctatus 

Plate  VI-4, 10 

dd.  Vitellus  1.8-1.4  mm.,  jelly  envelope  3.8  5.0  mm.  Season 
March  i  or  earlier  to  July  i  or  fall.  Hyla  arenicolor 

bb.  Envelope  1.1-1.0  mm. 

c.  Vitellus  0.6-0.8  mm.  Egg  complement  100.  Season  January  to 
September.  Pseudacns  ocularts 

Plates  VII  i ;  IX  34 

cc.  Vitellus  091.1  mm.  Egg  complement  809  1000.  Season  March 
30  to  May  15.  Hyla  c.  cructfcr 

Plates  VII-7,  IX  ii,LXVI  4 

BB.  Eggs  deposited  in  a  mass, 
a.  Egg  mass,  a  surface  film. 

b.  Single  egg  envelope  a  truncated  sphere,  the  flat  surface  above; 

egg  vitellus  black  above  and  white  below. 

c.  Envelope  1.8-4.0  mm.,  outline  always  distinct,  never  lost  in 

the  mass,  eggs  firm  and  distinct  like  glass  marbles,  making  a 

fine  mosaic;  vitellus  i .0-1.2.  mm.  Egg  complement  869.  Season 

March  to  Sept.  3.  Mtcrohyla  carohnensis 

Plates  V-6,7;  IX-56;  CXXV-5 


38  HANDBOOK  OF  FROGS  AND  TOADS 

cc.  Single  envelope  1.5-1.0  mm.,  vitellus  i.o  mm.  Egg  mass  a  loose  raft. 
Egg  complement  700.  Season  April  to  October.         Hyfofacbus  cuncus 

bb.  Egg  envelope  outline  indistinct,  more  or  less  merged  in  the  jelly  mass; 
jelly  glutinous,  envelope  not  a  truncated  sphere, 
c.  Egg  brown  above,  cream  or  yellow  below. 

d.  Egg  packets  small,  masses  seldom  if  ever  over  2.0  sq.  in.  (115  sq. 

cm.),  or  4  X  5  in.  in  diameter  (10  X  1^.5  cm.). 

e.  Inner  envelope  large,  1.1-3 .4  mm. ;  outer  envelope  3 .1-5  .o  mm. ; 

vitellus  0.8-1.6  mm.  Egg  complement  343-500.  Season  May  19 

to  Aug.  n.  Hyla  c.  cincrca 

Plates  VII  -9,  IX-io 

ee.  Inner  envelope  small,  1.4-1  o  mm.,  outer  envelope  4-8  mm. 
f.  Packets  small,  seldom  over  30-40  eggs;  vitellus  1.1-1.1  mm. 
Egg  complement  1803.  Season  May  10  to  Aug.  13. 

Hyla  v,  wrsicolor 
Plates  VII-n;  IX-i6;  LXXIII-5 

ff.  Packets  large,  sometimes  100-115  eggs,  vitellus  0.8-1.1  mm., 
av.  0.95  mm.  Egg  complement  708.  Season  April  i  to  Aug.  n. 

Hyla  femoralis 
Plates  VII -10,13;  IX-ii;  LXVIII-4 

cc.  Eggs  black  above  and  white  below. 

d.  Egg  mass  small;  outer  envelope  loose,  irregular,  does  not  look 
like  a  mosaic;  somewhat  merged,  inner  envelope  possibly  present 
close  to  vitellus,  vitellus  0.8-0.9  mm-  Outer  envelope  1.8-3.0 
mm.,  not  truncate.  Egg  complement  645.  Season  March  15  to 
September.  Mtcrohyla  oltvacca 

Plates  V-5;CXXVI-3 

dd.  Egg  packets  large,  loose,  glutinous  films,  35-675  sq.  in.  (118-3711 
sq.  cm.).  (Suspect  R.  heckschen  is  in  this  class.) 
e.  Inner  envelope  absent,  vitellus  1.1-1.7  mm.,  egg  mass  144-675 
sq  in  (900-3711  sq.  cm.)  in  area,  or  n  X  M  in.  (30  X  61  cm.) 
in  diameter,  egg  masses  among  brush  around  the  edge  of  ponds 
encircling  Pontca'cna-likc  vegetation  in  mid-pond.  Egg  comple- 
ment 10,000-10,000.  Season  March  15  to  Nov.  8. 

Rana  cattsbtiana 
Plates  VIII-7,  IX-45;  XCIV-6 

ec.  Inner  envelope  present,  1.8-4.0  mm.,  vitellus  1.4-1.0  mm. 

f.  Egg  mass  seldom  i  sq.  ft.  (35-144  sq.  in.  or  118-900  cm.) 
in  area,  or  5  X  7-1*.  in.  in  diameter;  usually  around  edge  of 
ponds,  inner  envelope  elliptic,  pynform,  or  circular,  av. 


GENERAL  ACCOUNT  39 

3.05  mm.;  vitellus  1.4-1.8  mm.,  mode  1.4  mm.,  av.  1.5 
mm.  Egg  complement  1451-4000.  Season  May  13  to 
Aug.  n.  Rana  clamttans 

Plates  VIII-9;  IX-4j;  XCV~4 

ff.  Egg  mass  over  i  sq.  ft.  in  area  (144-188  sq.  in.  or  900- 
1800  sq.  cm.),  or  n  X  n  in.  to  n  X  15  in.  in  diameter; 
usually  in  mid-pond,  inner  envelope  av.  3.45  mm.;  vitel- 
lus 1.4-1  o  mm.,  mode  1.8  mm.,  av.  17  mm.  Egg  comple- 
ment 8000-15,000.  Season  March  4  to  Sept.  15. 

Rana  ffylio 
Plates  VIIL  3,  IX-46 
aa.  Egg  mass  submerged, 
b.  Eggs  in  files  or  bands. 

c.  Eggs  laid  in  bands  which  soon  become  loose  cylinders  extending 
along  plant  stems  or  grass  blades. 

d.  Envelope  1.5-5.6  mm.,  vitellus  1.4-1.0  mm.,  vitelh  throughout 
jelly  cylinder,  if  stalkhke  on  almost  imperceptible  short  stalks. 
Egg  complements  1000-3000. 

e.  Envelope  3  8-5.6  mm.,  vitellus  1.4-10    Egg  complement 
1331.  Season  March  or  earlier  to  October. 

Scaphtopus  b.  bolbrookii 
Plates  V-i,4,IX-5,  XXI  5 

ee.  Envelope  1.5-3.5  mm.  Vitellus  1.4  1.6  mm.  Egg  comple- 
ment 1000-3000.  Season  April  to  August.    Scaphtopus  couchtt 

Plate  XVII  5 

dd.  Envelope  1.5  i.o  mm.,  vitellus  i  0-1.6,  eggs  in  bands  or  cylin- 
ders on  ends  of  grass  stems,  along  vegetation  stems,  etc.  Eggs 
near  periphery  of  cylinder  more  or  less  on  stalks  5  10  mm.  long. 
Egg  complement  1000-1000.  Season  mid-February  to  August. 

Scaphtopus  b.  bammondtt 
PlatcV-i,8,  XVIII  7 

cc.  Eggs  in  files. 

d.  Files  short  (4-10  mm.  in  length),  4-8  eggs  in  short  bcadhkc 
chain  or  bar,  or  many  such  files  radiating  from  one  focus,  vi- 
tellus 0.8-1.0  mm.,  tube  diameter  1.1  1.4  mm.  Egg  complement 
610,  766.  Season  April  i  to  Sept.  5.  Bufo  qucrctcus 

Plates  VI  i, 1,5,  IX- 14,  XL -5 

dd.  Files  long  (several  feet  in  length  or  often  a  meter  or  more  long), 
c.  Vitellus  i. 0-1.6  mm.,  tube  diameter  1.8-4.6  mm. 
f.  Inner  tube  absent. 


4o  HANDBOOK  OF  FROGS  AND  TOADS 

g.  Envelope  more  than  5  mm.  in  diameter  (5-6  mm.),  firm,  distinct;  vi- 
tellus  1. 1-1.6  mm.,  black  above,  white  or  gray  below;  35  eggs  in  30 
mm.  (i  3/16  in.).  Season  May  and  June.  Bufo  cahfornicus 

gg.  Envelope  less  than  5.0  mm.  in  diameter  (1.8-4.6  mm.). 

h.  Envelope  i  6-4.6  mm.,  distinct  and  firm;  vitelli  crowded  in  the 
files  at  first  in  double  row,  later  more  spread  out  but  still  crowded 
although  at  times  in  single  row,  11  15  eggs  in  30  mm.  (i  3/16 
in.).  Season  March  to  May.  B.  woodhousii 

i.  Egg  complement  smaller,  5000-10,000;  egg  envelopes  possibly 
smaller.  Eastern  Missouri  eastward.  Season  April  to  mid- August. 

B,  w.  fowleri 
Plate  IX-i7a,b 

11.  Egg  complement  larger  to  15,000,  western  Missouri  westward. 
Season  March  to  July.  B  w.  woodhousii 

hh    Envelopes  i  4  or  less  (i  8-1.4  mm  ) 

i.  Envelope  slightly  scalloped  in  appearance;  tube  tightly  coiled; 
1.8  1.4  mm.,  narrow,  vitelli  crowded,  vitelh  grayish  or  greenish 
brown  above,  cartridge  buff  or  sulphur  yellow  below,  14-30 
eggs  in  30  mm.,  vitcllus  i  1-1.6  mm.  Season  May  i  to  July  10. 

Bufo  compacttlts 
Plates  VI-9,  XXXIII-4 

li.  Envelope  not  scalloped,  rather  loose  but  distinct  in  outline, 
vitelli  black  or  deep  brown  above,  tan  or  white  below,  i.r- 
1.7  mm  in  diameter  Season  March  to  Aug  15.  Egg  comple- 
ment 7500-8000.  Bufo  alvanus 

ff .  Inner  and  outer  tubes  present  (i  in  number), 
g.  Partitions  apparent  between  eggs. 

h.  Outer  envelope  distinctly  scalloped;  envelopes  laminated,  outer  en- 
velope 1.7  mm.  at  emargmation,  i.o  mm.  at  widest  diameter  if  in 
single  row,  or  i.o  mm.  and  1.6  mm.  if  eggs  in  double  row;  inner 
envelope  1.6  mm.  Vitellus  1.1  mm  ,  black  above,  white  below, 
complement  to  10,000.  Season  April  to  September.  B.  cognatus 

hh.  Outer  envelope  not  scalloped,  3  4-4.0  mm.  in  diameter,  usually  not 
two  rows,  inner  tube  distinct  1.6-1.1  mm.;  vitellus  1.0-1.4  mm. 
black  or  dark  brown  above,  white  below,  18-10  eggs  in  30  mm. 
(i  3/16  in.).  Egg  complement  4000-10,000.  Season  March  to  late 
July.  Bufo  a.  amencanus 

Plates  VI-7,  IX-8;  XXV-6 


GENERAL  ACCOUNT  41 

gg.  No  partitions  between  each  individual  egg. 

h.  Eggs  in  a  single  row  within  the  jelly  tube.  Outer  en- 
velope 1.6-4.6  mm.;  inner  tube  close  to  the  outer 
tube,  inner  tube  1.1-3.4  mm.;  outer  tube  inclined  to 
be  slightly  scalloped,  distinct  space  between  eggs; 
vitelli  i. 3-1. 4  mm.;  7-8  eggs  in  30  mm.  (i  3/16  in.). 
Season  last  of  February  to  September.  Bufo  temstrts 

Plates  VI-6;IX-i 5 

hh.  Eggs  usually  in  a  double  row  within  jelly  tube, 
more  crowded. 

i.  Vitelli  small,  1.1  mm.,  outer  envelope  small,  1.8  - 
3.1  mm.,  purplish  black  and  white,  inner  envelope 
1.6  mm.,  close  to  outer  tube,  10  eggs  in  30  mm. 
(single  row),  or  15-17  eggs  in  30  mm.  (double 
row),  rarely  single  eggs.  Season  March  to  Aug.  15. 

Bufo  valltceps 
Plate  VI  3,8 

ii.  Vitellus  larger,  1.50-1  7  mm  ,  black  and  white  or 
cream,  outer  envelope  loose  but  distinct,  large, 
4  9-5.3  mm.  in  diameter,  vitelli  double-  or  triple- 
rowed,  inner  tube  3.5-3.8  mm  ,  not  very  close  to 
outer  tube  (boreas  group). 

j.  Vitelli  larger,  1.65-1.75  mm   Season  January  to 
July.  Bufo  boreas  halophtlus 

)j.  Same,  except  vitelli  average  smaller  though 
their  range  is  1.5-1.75  mm.  Season  March  to 
September.  Bufo  b.  boreas 

bb.  Eggs  an  irregular  mas>s,  jelly  loose,  envelope  3.1  6.7  mm.,  vitellus 
black  above  and  white  below. 

c.  Single  envelope  3.1-3.6  mm.,  vitellus  j  a  1.3  mm  ,  egg  mass  sub- 
merged film  on  bottom  or  loosely  strewn  on  bottom.  Season  April 
to  September.  Bufo  punctatus 

Plate  VI-  4,10 

cc.  Single  envelope  6.3  mm.,  vitellus  1.3  mm.,  egg  mass  various  pat- 
terns, single,  tandemlike,  Bufo-hkc,  lattice  work,  etc.,  at  or  near 
surface  attached  to  vegetation.  Season  April  to  June. 

Scaphiopus  h.  hurt  mi 


42  HANDBOOK  OF  FROGS  AND  TOADS 

bbb.  Eggs  in  lumps. 

c.  Egg  mass  a  firm  regular  cluster. 

d.  One  jelly  envelope.  (At  times  R.  syhatica,  R.  s.cantabngtnsis,nn& 
R.  p.  pntiosa  appear  in  this  category.) 

e.  Egg  mass  a  plinth  or  sometimes  globular,  firm.  No  indistinct 
inner  envelope. 

f.  Vitellus  1.4-1.8  mm.,  black  above,  sulphur  or  primrose 
yellow  below,  envelope  4.9-6.9  mm.,  av.  5.4  mm.,  eggs 
farther  apart  than  in  R.  ptpicns  or  R.  p.  sphcnocephala;  egg 
complement  350-500  eggs,  egg  mass  1.5  X  i  to  3  X  4  in. 
New  Jersey,  southward.  Season  June  11  to  Aug.  n. 

Rana  virgatipes 
Plates  VHI-i,IX-55 

ff.  Vitellus  1.8  11  mm.,  black  above,  light  tan  below;  en- 
velope 5-71  mm.,  av.  63,  egg  complement  1000-3000 
eggs  Egg  mass  3  x  3  to  8  x  6  in.  Northern  Nevada  north- 
ward and  westward.  Season  March  to  May. 

R.  prctiosa  lutcivcntns 
Plate  VIII -6 

ec.  Egg  mass  a  sphere,  inner  envelope  indistinct.  (See  dd.  Two 
or  three  jelly  envelopes  R  s  syhattca,  R.  s.  cantabrtgensts,  R. 
p.  prettosa.} 

dd.  More  than  one  jelly  envelope, 
c.  Two  envelopes. 

f.  Egg  mass  a  sphere  1^-4  in.  (6  35-10  cm.)  in  diameter,  con- 
taming  1000-3000  eggs,  outer  envelope  distinct, 
g    Eggs  black  above  and  white  below,  inner  envelope  ap- 
parently absent,  slightly  evident  under  lens,  3.6-5.8 
mm.,  av.  3.8,  outer  envelope  5.1-9.4  mm.,  av.  6.4,  vi- 
tellus 1.8-1  4  mm.,  av.  1.9.  Egg  complement  1000-3000. 
Season  March  19  to  May  i.  Rana  s.  sylvattca 

Plates  VIII-8,  IX-53,  CXIX~5 

[h.  Is  R.  s  cantabngcnsis  thus:  vitellus  1.50-1.8  mm.,  av. 
i  65,  inner  envelope  3-4  mm  ,  av.  3.5  mm.,  outer 
envelope  4.1-5.4  mm.,  av.  5  mm.?] 

gg.  Eggs  brown  above  and  yellow  below,  inner  envelope 
distinct,  13-3.0  mm,  outer  envelope  3.6-5.0  mm.; 
vitellus  1.6-1.9  mm.  Egg  complement  1000  to  3000. 
Season  April  6  to  May  18.  Rana  palustris 

Plates  VIII-4;  1X^47 


GENERAL  ACCOUNT  43 

ff.  Egg  mass  a  plinth,  complement  large,  1000-10,000  eggs;  eggs  black  above 
and  white  below. 

g.  Outer  envelope  large,  10-15  nim.,  inner  envelope  indistinct,  some- 
times absent,  5.0  6.0  mm.,  vitellus  x-x.8  mm.,  black  and  white,  far 
apart  in  mass  like  R.  syhatica;  egg  complement  1000-1000  eggs  in 
masses  as  large  as  8  X  6  in  Northwest.  Season  March  to  May. 

Rana  p.  prctiota 

gg    Outer  envelope  smaller,  7  o  mm.  or  less,  inner  envelope  i  4-4.0  mm., 
vitelli  i.4~x.5  mm  ,  black  and  white,  close  together  like  R   p/p/wr 
masses  Except  for  R.  p/p/cns,  east  of  Rockies, 
h.  Vitellus  av  xa-x.4orx5mm 

i.  Vitellus  av.  x  5  (range  x  4  x  6  mm.).  Outer  envelope  4.5-5.0 
mm  ,  distinct,  inner  envelope  3.x  mm.  Masses  larger,  5000 
10,000  eggs   Midwest.  Season  March  to  May  15. 

Rana  arcolata  circulosa 

li.  Vitellus  av.  x  o  mm   (range  18x4  mm  ),  inner  envelope  3.1 
4  4  mm  ,  outer  envelope  4460  mm  ,  mode  5  x  mm.,  av.  5.3 
mm.  Egg  complement  5000  or  more   Southeast.  Season  March 
13  to  Nov   3  Ram  captto 

Plates  VIII  x,  IX  44,  XV  i,  x,  XCII  4 

hh.  Vitellus  av  1.4  i  7  mm.  (range  13x0  mm  ),  inner  envelope  x  3  - 
3  x  mm  ,  outer  envelope  3466  mm.  Egg  complement  1000  5000. 
i.  Av.  outer  envelope  5  o  mm  or  larger. 

)    Av.  outer  envelope  6  3  mm.  (5666  mm  ),  inner  envelope 
x  4  3  o  mm  ,  av.  x  75  mm.,  vitellus  j  4  av.,  range  1.3  i  6 
mm  ,  vitellus  black  or  brown  above,  white  or  yellow  below 
Egg  complement  800  1500.  Season  June  X5  to  July  30 

Rana  tcpttntnonahs 
Plate  CXVIII  3 

jj.  Av.  outer  envelope  5  i  mm  (range  4x  6.0  mm.,  mode  5.0 
mm  ),  inner  envelope  1534  mm.,  av.  x  X5  mm  ,  vitellus 
j  7  mm.,  range  i  3  x.o  mm.  Egg  complement  3500  4500. 
Season  March  30  to  May  15  Rana  p.  piptws 

Plate  1X^48,0-3, 4 

jjj  Av.  outer  envelope  3  8  mm  (range  3454  mm  ,  mode  4  o 
mm  ),  inner  envelope  x  4  3  x  mm  ,  vitellus  i  4  1.8  mm., 
av.  1.6  mm.,  egg  complement  1084.  Season  January  to  De- 
cember. Rana  p.  sphcnoccphala 

Plate  IX-49 


44  HANDBOOK  OF  FROGS  AND  TOADS 

cc.  Three  envelopes. 

f.  Outer  envelope  3.8-4.5  mm.,  av.  40  mm.;  firm;  middle  envelope 
1.6-3.4  mm.,  av.  1.8  mm.;  inner  envelope  1.3-3.0  mm.,  av.  1.5  mm. 
All  envelopes  distinct.  Vitellus  1.9-1.5  mm.,  av.  1.1  mm.,  black  and 
white.  Mass  compact  i  X  2.  X  1.1  in.  to  i  X  4  X  M  in.  Eggs 
firmly  attached  to  each  other.  Egg  complement  900-1100.  Season 
March  I  to  May  i.  Rana  b.  boylti 

Plate  VIII -5 


ff.  Outer  envelope  6.4-14.0  mm. 

g.  Egg  mass  small,  400-500  or  less,  all  envelopes  distinct. 

h.  Outer  envelope  6.4-7.9  mm.,  av  7.1  mm.;  middle  envelope 
4.1-5.0  mm.,  av.  4.6  mm.,  inner  envelope  1.75-4.8  mm.,  av. 
3.9  mm.,  vitellus  1.8-1.3  mm.,  av.  1.1  mm.,  black  above  and 
light  gray  tan  below,  phnthhke  mass  18  X  40  mm.  with  eggs 
J^-J^  in.  apart.  Egg  complement  100-400  eggs.  High  Sierras. 
Season  June,  July.  Rana  b.  stcrrae 

hh.  Outer  envelope  1.1-1.1  mm.,  middle  envelope  5-6  mm.,  inner 
envelope  4.9  mm.,  vitellus  1.15  mm  ,  black  above,  cream  be- 
low. Mass  400-415  eggs.  Washington  and  northern  Oregon. 
Season  last  of  May  to  July  i.  Rana  cascadae 

gg.  Egg  mass  large,  750-4000.  Envelopes  quite  distinct  or  indistinct, 
h.  All  envelopes  indistinct,  especially  the  middle  one.  Outer 
envelope  10-14.0  mm.,  middle  envelope  6.1-8.0  mm.,  av.  6.8 
mm.,  inner  envelope  40-6.7  mm.,  av.  5.7  mm  ,  vitellus  1.3- 
3.6  mm.,  av.  3.0  mm  ,  black  above,  creamy  white  below. 
750-1400  eggs  in  a  flat  mass  6  X  10  in.  across,  vitelh  *±  in. 
apart.  Principally  northwest  Oregon  northward.  Season  Feb- 
ruary to  May.  Rana  a.  aurora 

hh.  All  envelopes  quite  distinct,  middle  one  particularly  dense 
and  distinct,  outer  envelope  7.5-11.8  mm.,  av.  8.5  mm.,  mid- 
dle envelope  3.9  6.4  mm.,  av.  4.40  mm.,  inner  envelope  3.9-- 
6.4  mm.,  av.  4.4  mm.  Vitellus  1.0-1.8  mm.,  av.  1.1  mm.; 
black  and  creamy  white,  egg  complement  1000-4000  eggs 
in  a  soft,  viscid  mass  1.6  X  4  X  3  inches  to  6  X  4  X  4  in. 
Principally  California  and  Lower  California.  Season  January 
to  March.  Rana  a.  draytonri 


GENERAL  ACCOUNT  45 

cc.  Egg  mass  a  loose  irregular  cluster,  sediment-covered. 

d.  Egg  mass  small,  generally  about  or  less  than  i  in.  (1.5-3.5  cmO  m 
diameter,  normally  10-15  or  4°»  r*rely  100  or  more  in  one  mass, 
e.  No  inner  envelope. 

f.  Smaller  species  19-37.5  mm.  (19-31  males,  19-37.5  females). 
Egg  or  vitellus  0.615-1.1  mm.,  av.  below  1.1  mm.  (except  in 
P.  «.  septentrtonalts,  1.1-1.4  mm.)  single  envelope  1.6-5.0  mm. 
normally,  rarely  to  7.8  mm. 

g.  Vitelh  1.1-1.4,  envelope  larger.  Season  May  and  early  June. 

Pseudacris  nigrita  septentrionalts 

gg.  Vitelh   0.615-1.1  mm.;   envelope   1.6-5.0  mm.    normally. 
Season  early  February  or  earlier  to  May. 
h.  Envelope  1.1  1.4  mm.  or  more,  vitellus  0.615-  9  mm. 
Season  March  10  to  May  10.  Pseudacris  n.  darkii 

Plates  VII-4,  L~4 

hh.  Envelope  i  6-1.8,  vitellus  0.9  i.o  mm.  Florida  to  Georgia. 
Season  February  to  August  15.  Pseudacrts  ;/.  verrucosa 

hhh.  Envelope  3.0-5.0,  sometimes  more. 

i.  Envelope  3  1-4.0  mm.,  vitellus  0911  mm.  Vitellus 
black  and  cream  or  slightly  yellowish.  Season  February 
to  May.  Pseudacrts  n.  fertarum 

Plates  VII  1,5,  LI  4 

ii.  Envelope  3.0-  5.9,  sometimes  7  8  mm.,  vitellus  0.9  1.1 

mm.  Egg  complement  500  800    Season  March  19  to 

May.  Pseudacrts  n.  truer  tat  a 

Plates  VII-6,  IX  }ia,b,c,  LIII  4 

ff.  Larger  species  14  48  mm.  (14  41  males,  17-48  mm.  females). 

Egg  or  vitellus  av.  over  i  i  mm.  (range  1.1-2.0  mm  ).  Single 

envelope  3.0-8  5  mm.  Masses  IQ  100,  rarely  100  in  a  mass. 

g.  Envelope  6.0  8.5  mm.,  vitellus  1.6  mm.  West  Virginia  to 

Ohio  to  northern  Mississippi  and  Alabama.  Season  March 

to  July.  Pseudacris  brachyphona 

gg.  Envelope  3.0-7.15  mm.,  av.  normally  under  6.0  mm. 

h.  Vitelh  1.6  mm.  Envelope  3.1-4.1  mm.,  av.  under  4.0  mm. 
Southeast.  Season  November  to  March.   Pseudacris  ornata 


46  HANDBOOK  OF  FROGS  AND  TOADS 

hh.  Vitclh  i.z-i.3  mm.  (Bragg;  1.1-1.8  or  10,  Strecker's  ma- 
terial), envelope  3.0-7.15  mm  ,  av.  over  5.0  mm.  Texas  to 
Oklahoma.  Season  December  to  late  May. 

Pstudacris  strtckcri 
ee.  Inner  envelope  i  4  1.7  mm. 

f.  Outer  envelope  smaller,  1.1-3.0  mm.,  vitellus  smaller,  0.65-1.1 
mm.,  inner  envelope  1.4-1.1  mm. 

g.  Outer  envelope  small,  1.1-1  4  mm.  or  more;  inner  envelope 
1.4  1.8  mm.,  vitellus  0.65-0.9  mm  ,  mass  smaller,  13-30  eggs; 
egg  complement,  150-300  eggs.  Season  March  5  to  mid-August. 

Pscudacns  n.  clarkti 
Plate  VII-4,  L-4 

gg.  Outer  envelope  larger,  3.0-5  o  or  7.8  mm.,  inner  envelope  1.1 
mm.;  vitellus  o  91  i  mm  Mass  15-300  eggs  per  mass.  Egg 
complement  500  -1500  eggs.  Season  March  10  to  May  10. 

Pseudacrn  n.  tnsmata 
Plates  VII-6,  IX  3ia,b,c,  LIII-4 

ff  Outer  envelope  larger  4.7-6  7  mm.,  inner  envelope  i  8-1.7  mm.; 
vitellus  larger  13135  mm  ,  vitellus  light  brown  above  and  pale 
white,  yellowish,  or  greenish  below.  Masses  9-75  eggs,  rarely 
single.  Egg  complement  600- 1500.  Hyla  regilla 

dd.  Egg  mass  an  irregular  cylinder  1-6  in  (i  5-15  cm.)  in  length,  extending 
along  plant  stem  or  grass  blade,  envelope  single,  3.8-5  6  mm  ,  vitellus 
i  .4  -i.o  mm.  Egg  complement  1331.  Season  April  or  earlier  to  August  17. 
(See  also  d,  dd,  c,  cc  under  c  Eggs  laid  in  bands  which  soon  become 
loose  cylinders  extending  along  plant  stems  or  grass  blades  ) 

Scapbtopus  fy.  bolbrooktt 
Plates  ¥-1,4,  IX-5,XXI-5 

We  never  described  in  detail  our  results  from  work  in  Texas  during  1925- 
19^5.  The  Florida  group  of  Carr,  Gom,  and  Van  Hynmg  have  added  much 
to  our  knowledge  of  the  Florida  forms.  H.  M.  Smith  in  1934  did  a  good  study 
of  the  Kansas  forms.  J.  R.  Slater  of  Tacoma  has  described  the  forms  of  the 
Northwest  (still  in  manuscript),  and  A.  and  R.  D.  Svihla  and  K.  Gordon 
have  also  added  data  about  that  area.  But  the  group  which  has  been  most 
vigorous  is  the  Oklahoma  group  (from  the  University  of  Oklahoma — Bragg, 
Trowbridge,  Ortenburger,  and  others;  and  from  Oklahoma  A.  and  M.  Col- 
lege, the  Moore  group). 

Tadpoles  (Plates  X-XV:  lateral  views,  X-XI;  mouth  parts,  XII-XIV;  de- 
velopment, XV) :  Some  25  to  30  species  of  tadpoles  of  the  United  States  and 


GENERAL  ACCOUNT  47 

Canada  need  to  be  described.  Most  have  been  found  but  were  not  described, 
and  the  tadpoles  of  15  forms  are  yet  unknown  to  science.  The  life  histories 
of  the  robber  frogs  we  know  in  our  country  indicate  no  free  tadpole  stage, 
the  whole  development  being  in  the  egg. 

In  discussions  of  the  size  of  tadpoles  quite  small  is  i  inch  (24  mm.)  or 
smaller;  small,  T-I%  inches  (24-3*5  mm.) :  medium,  i3£-2  inches  (40-50  mm.) ; 
large  2%-$%  inches  (60-86  mm.);  quite  large,  ?/.:>-4  inches  (95-100  mm.); 
very  large,  5%-5  Jfi  inches  (135-145  mm.).  Some  of  the  toads  and  swamp 
cricket  frogs  may  have  tadpoles  a  little  less  than  i  inch  (23  or  24  mm.)  in 
length,  whereas  bullfrogs  may  have  tadpoles  5%  inches  (145  mm.)  or  even 
larger. 

The  tadpole  has  a  body  and  a  tail.  The  body  has  sensory  lines,  a  breathing 
pore  or  spiracle,  a  vent  or  anus,  eyes,  nostrils,  and  a  mouth.  The  mouth  is 
wholly  unlike  the  adult  mouth.  It  usually  has  a  disk  called  a  Libium  (with 
upper  and  lower  labia).  Generally  about  the  edge  of  the  labium  are  tubercles 
or  papillae.  At  the  inner  edge  of  each  labium  or  at  the  very  portal  of  the  mouth 
opening  itself  are  horny  crescents  called  upper  and  lower  mandibles.  On  the 
upper  and  lower  labia  arc  horny  ridges  of  teeth  or  combs  for  scraping  food. 
The  tail  has  two  parts:  the  axis  consists  of  muscle  segments;  and  the  fin  con- 
sists of  upper  and  lower  crests. 

The  narrow-mouthed  toad  tadpole  has  no  mouth  disk,  no  labial  teeth,  no 
papillae,  no  horny  beak.  The  spiracle  is  next  to  the  vent.  The  eyes  are  on  a 
lateral  ndgc.  These  tadpoles  are  small  black  flattened  creatures  with  some 
white  on  the  tail  axis  or  body. 

The  ribbed  toad  (Ascaphus)  has  a  mouth  disk,  an  upper  mandible  only, 
papillae  on  the  lower  lip  or  labium,  upper  labial  teeth  at  least  two  rows  to  a 
ridge,  labial  tooth  ridges  two  to  three  above  and  seven  to  ten  below,  a  spiracle 
in  the  middle  of  the  venter  nearer  the  hind  legs  than  the  snout,  and  eyes 
straight  back  of  and  close  to  nostrils,  dorsal,  and  equidistant  from  micldorsal 
line  and  lateral  outline  when  viewed  from  above.  The  tail  is  spatulatc  and 
rounded  as  in  some  mountain  stream  tadpoles.  The  tadpoles  are  black  or 
brown  with  white  tips. 

The  spadefoots  have  the  vent  in  the  middle  position,  the  spiracle  below  the 
body  axis  and  on  the  left  side,  the  papillae  completely  around  the  labium 
except  for  a  small  interval  above  (absent  in  one  species),  the  papillary  border 
not  emargmate  on  the  side,  labial  teeth  three  to  six  ndges  above  and  four  to 
six  ridges  below,  eyes  dorsal  and  nearer  middorsal  line  than  lateral  outline, 
and  muscle  segments  of  the  tail  plainly  visible. 

The  toads  usually  have  small  blackish  tadpoles  with  anus  median,  papillae 
confined  to  the  sides  of  the  labium,  upper  and  lower  edges  of  the  labium 
toothed,  sides  of  labium  emarginate,  labial  teeth  two  ridges  above  and  three 
ridges  below,  and  eyes  dorsal  and  slightly  nearer  the  lateral  outline  than  mid- 
dorsal  line.  The  spiracle  is  on  the  left  side  and  is  small. 


;;;,;:, :;; 


Plate  X.  Mature  tadpoles  (X1)-  i-  Anderson  tree  frog,  Hyla  andcrsomt,  2.  Green  tree  frog,  Hyla 
cinerca.  3.  Peeper,  Hyla  cmctfct  cntcifer.  4.  Pine  woods  tree  frog,  Hyla  jcmonihs.  5,6.  Barker, 
Hyla  gratiosa.  7.  Squirrel  tree  frog,  Hyla  Stjtwella.  8.  Little  chorus  frog,  Pscudacns  octdans.  9,10. 
Common  tree  toad,  Hyla  vcrsicolor  vcrsicohr.  u.  Cricket  frog,  Acns  gryllns  gryllus.  12.  Spadefoot, 
St'aphiopus  holbrookti  holbroofyi.  13.  Oak  toad,  Bttfo  qttercictts.  14.  Narrow-mouth  toad,  Micro- 
hyla  carolincnsis.  (Wright,  Gen.,  1932,  pp.  46-47.) 


;,^w:^  vf.' 


13 


X/.  /?«««  tadpoles  (mature  XO-  1»2-  Gopher  frog,  /?.  r0/7/'/0.  3,4.  Green  frog,  /^.  damitans. 
5,6.  Southern  bullfrog,  K.  grylto.  7,8.  River-swamp  frog,  /?.  hec^schcn.  9,10.  Southern  meadow  frog, 
R.  piptcns  sphenocephala  11.  Mink  frog,  R.  septentnonahs.  12.  Meadow  frog,  R.  p.  pipiens.  13. 
Sphagnum  frog,  R.  virgatipcs.  14.  Pickerel  frog,  R.  palustris.  (Wright,  Gen.,  1932,  pp.  45-47.) 


rlatc  XII.  Tadpole  mouth  parts  (Hyla,  Acris,  Pseud acris,  Bttfo).  i.  Peeper,  H.  crucijer  crucijer. 
2.  Barker,  H.  gratiosa.  3.  Anderson  tree  frog,  H.  andersonn.  4.  Green  tree  frog,  H.  cinerea.  5.  Eastern 
swamp  cricket  frog,  P.  nigrita  jcnarum.  6.  Cricket  frog,  A.  gryttus  gryllus.  7.  Fowler's  toad,  B. 
woodhousii  jotilcn.  8.  Little  chorus  frog,  /'.  octtlans.  9.  Squirrel  tree  frog,  H.  sqmrella.  10.  American 
toad,  B.  amencamts  americanus.  n.  Southern  toad,  B.  tetrestris.  12.  Common  tree  toad,  H.  vcrsicolor 
versicolor.  13.  Pine  woods  tree  frog,  H.  fcmorahs.  (Wright,  Gen.,  1932,  pp.  54"55-) 


Plate  Kill  Tadpole  mouth  parts  (Rana,  Hyla,  Pscudacrts,  Bu)o,  Ascapftus).  i.  spnagnum  irog, 
R.  virgatipes.  2.  Nevada  frog,  R.  fisheri.  3.  Canyon  tree  frog,  H.  arenicohr.  4.  Clarke's  chorus  frog, 
P  nignta  claret.  5.  Spotted  toad,  B.  punctatus.  6.  Spadefoot  toad,  B.  compactihs.  7.  Nehulous  toad, 
B.  valliccps.  (Wright  and  Wright,  MS,  1930.)  8.  American  bell  toad,  A.  truci.  (After  Gaige,) 


52  HANDBOOK  OF  FROGS  AND  TOADS 

The  frogs  have  medium-to-large  tadpoles,  with  spiracle  on  left  side,  vent  on 
right  side  of  lower  tail  fin's  base.  The  spiracle  is  near  the  body  axis;  the  papil- 
lary border  is  cmarginatc  on  the  side;  the  labial  teeth  are  two  or  three  ridges 
or  more  above  and  three  or  four  or  more  below.  Some  may  have  spatulate 
tails,  if  high  mountain  forms,  others  may  have  upper  tail  crest  high  and  far 
on  the  body  like  a  tree  frog  tadpole,  but  most  have  the  tail  crests  neither  very 
high  nor  extending  far  on  the  body. 

The  tree  frogs  have  small  or  medium  tadpoles,  often  with  high  dorsal  tail 
crest  extending  far  on  the  body,  the  vent  on  right  side  of  ventral  tail  crest,  and 
the  spiracle  on  left  side  near  the  body  axis.  The  labial  teeth  have  two  ridges 
above  and  two  or  three  below. 

The  following  synopsis  of  tadpoles  is  offered  only  after  considerable  de- 
bate. We  could  have  put  it  in  the  first  edition.  No  one  knows  the  extent  of 
variation  in  the  mouth-part  characters.  If  one  attempts  to  use  other  workers' 
results,  he  encounters  personal  equational  differences  in  measuring  technique. 
We  might  find  no  median  space  in  the  second  upper  lateral  tooth  row  in 
some  P.  n.  clarion  and  a  median  space  in  others.  Dr.  Bragg  figures  his  material 
with  no  median  space,  as  we  at  times  have  found.  What  is  normal  for  the 
whole  range  of  a  species?  This  synopsis  is  presented  in  the  hope  that  some- 
one will  attempt  to  learn  the  frog  tadpoles  of  the  United  States  in  the  field 
and  refine  our  descriptions.  Grace  L.  Orton  may. 


Plate  XIV.  Tadpole  mouth  parts  (Rana,  Mtcrohyla,  Scaphiopus).  i.  River-swamp  frog,  R. 
hecfahen.  2.  Young  of  (i).  3.  Mink  frog,  R.  scptcntrwnahs.  4.  Sphagnum  frog.  R.  virgatipcs.  5. 
Southern  bullfrog,  R.  grylto.  6.  Narrow-mouth  toad,  M.  carohncnsis.  7.  Green  frog,  R  cl-""""f 
8.  Spadefoot,  S.  holbrootyt.  (Wright,  Gen.,  1932,  pp.  54-55.) 


§4  HANDBOOK  OF  FROGS  AND  TOADS 

SYNOPSIS  OF  UNITED  STATES  TADPOLES 

A.  No  tadpole  stage. 

Eleutherodactylus  ricordn  plantrostns  (Plate  LXXVIII-5,7,8),  E.  latrans; 
Syrrhophus  camp;  S.  gatgeae;  S.  marnockii. 
AA.  A  tadpole  stage  in  water. 
B.  Tadpoles  unknown. 

Scaphiopus  holbrookii  albus;  Bufo  caltfornicus;  B.  hemiophrys;  Pseudacris 
n.  nigrtta;  P.  occidental**;  P.  n.  septentrional ts;  Hyla  baudtnii;  H.  v. 
phatocrypta;  H.  avwoca;  H.  vcrsicolor  chrysosccltsj  Rana  a.  aurora;  R. 
prettosa;  Mtcrohyla  anolata;  Hypopachus  cuneus — 14  in  all. 
BB.  Tadpoles  seen  but  not  described  in  detail. 

Bufo  alvarius;  B.  b.  boreas;  B.  canorusj  B  debths;  B.  woodhousii  fowleri; 
Pseudacris  ornata;  Hyla  cinerea  cvittata;  H.  wrtghtorum;  Rana  a.  areolata; 
R.  sylvattca  cantabrtgensis;  R.  fisher*. 
BBB.  Tadpoles  in  this  synopsis. 

a.  Mouth  disk  absent,  no  labial  teeth;  no  papillae,  no  horny  beaks; 
spiracle  median  near  anus;  nostril  within  edge  of  mouth  fold,  eye  on  a 
canthus;  tadpoles  (2.3-16  4  mm.)  small,  black  or  grayish  olive  tadpoles 
with  a  stripe  on  the  middle  of  the  tail  musculature. 

BREVICIPITIDAE 

b.  Tail  tip  always  black,  eyes  just  visible  from  ventral  aspect;  black 
pointed  excrescences  back  of  upper  labial  edge,  lower  mandibular 
prolongation  gray,  coloration  citrine-drab  or  grayish  olive,  with 
middorsum  dark  grayish  olive,  venter  with  white  or  pale  pinkish 
cinnamon  spots,  sides  of  body  without  striking  longitudinal  light 
bands,  light  band  at  base  of  tail  musculature  not  prominent  (in 
alcohol).  Outer  egg  envelope  not  truncate,  mass  seldom  showing 
distinct  outline  of  each  egg  envelope  in  a  mosaic  fashion.  Texas 
westward  to  Fort  Davis  Mountains.  Mtcrohyla  olwacea 

Plate  CXXVI-4,5 

bb.  Tail,  sometimes  with  black  tip,  eyes  plainly  visible  from  ventral 
aspect,  inner  face  of  upper  labial  edge  with  no  black  pointed  ex- 
crescences; lower  mandibular  prolongation  light,  general  coloration 
black  with  purplish  gray  or  hair  brown  dots,  venter  with  white 
or  yellowish  bands  and  large  blotches,  sides  of  body  with  same 
coloration;  light  band  at  base  of  tail  musculature  prominent  (even 
in  alcohol).  Egg  with  truncate  outer  envelope,  giving  mass  a  mosaic 
appearance  on  water's  surface.  Virginia  to  Florida  to  Texas  and  up 
Mississippi  River  to  Indiana.  Mtcrohyla  caroltnensts 

Plates  X-i4,  XIV-6;  CXXV-4 


GENERAL  ACCOUNT  55 

aa.  Mouth  disk  present;  upper  and  lower  labial  teeth,  labial  maxillae;  at 
least  the  upper  horny  beak  present,  nostril  free  of  mouth, 
b.  Spiracle  median  nearer  insertion  of  hind  legs  than  tip  of  snout;  labial 
teeth  1/7  to  3/10,  upper  labial  teeth  at  least  two  rows  to  a  ridge;  no 
lower  beak;  many  rows  of  papillae  on  edge  of  lower  labium,  mouth 
large  and  round;  anus  median,  black  or  blackish  brown  tadpoles 
speckled  with  black;  tail  may  be  of  body  color  or  spotted  with 
creamy  white,  tail  tip  white  or  yellowish  white;  upper  tail  crest  not 
extending  on  the  body.  Washington  to  California. 

Ascaphus  truet  ASCAPHIDAE 
Plates  XIII-8,  XVI-i 

bb.  Spiracle  sinistral,  upper  labial  teeth  not  with  two  close  rows  on  each 
ridge,  labial  teeth  1/3,  1/1,  1/3,  1/4,  3/3,  3/4,  4/4  to  6/5,  upper 
and  lower  horny  beaks;  papillae  on  lower  edge  of  labium  absent  or 
in  one  or  two  rows. 

c.  Anus  median,  spiracle  lateral  below  the  lateral  axis  (of  tail  mus- 
culature, projecting)  sometimes  as  much  ventral  as  lateral,  upper 
tail  crest  extends  on  the  body  to  a  vertical  nearer  hind  legs  than 
the  spiracle  or  only  halfway;  viscera  visible  (in  preserved  speci- 
mens) through  the  skin  of  the  belly.  Eyes  distinctly  dorsal, 
d.  Labial  teeth  3/4  to  6/6,  papillae  extending  completely  around 
the  border  of  the  labium  except  for  a  short-toothed  median 
interval  above  (sometimes  absent  in  one  species);  papillary 
border  not  emarginate  on  each  side,  eyes  nearer  middorsal  line 
than  lateral  outline,  on  lateral  axis,  tadpoles  14.5-65  mm.  in 
length,  spiracle  a  slit,  very  low  on  side,  about  on  the  level 
of  the  mouth,  in  general  very  bronzy  tadpoles,  myotomes  of 
tail  musculature  well  indicated.  Tail  tip  usually  rounded. 

SCAPHIOPODIDAE 

e.  Teeth  6/6,  5/6,  6/5,  5/5,  tadpoles  small,  2.1-18  mm.,  trans- 
forming at  8.5-11.0  mm.,  inner  papillae  present,  spiracle 
equidistant  between  eye  and  base  of  hind  legs  or  vent,  eye 
1.4-1  8  nearer  tip  of  snout  to  the  spiracle,  av.  1.51,  inter- 
nasal  space  in  interorbital  space  1.18-1.83,  av-  I-5^J  depth 
of  body  1.75-1.5  in  body  length,  av.  1.04,  muscular  part 
of  tail  in  depth  of  tail  1.45-1.5,  av.  1.98,  last  lower  row 
of  teeth  longer  than  horny  beak.  Last  lower  row  of  teeth 
1.5  times  in  next  to  lowest  row  of  teeth.  Egg  mass  an  ir- 
regular cylinder,  at  first  bandhke.  Massachusetts  to  Florida 
to  Texas  and  Arkansas.  Scaphiopus  h.  holbrookii 

Plates  X  -ii,  XIV-8 


56  HANDBOOK  OF  FROGS  AND  TOADS 

cc    Labial  teeth  normally  4/5  or  4/4,  sometimes  5/4,  3/4,  or  possibly  1/4; 

inner  papillae  generally  absent,  tadpoles  small  or  large;  eye  1.0-1.6 

nearer  tip  of  snout  than  spiracle;  last  lower  row  of  teeth  less  than  or 

J^  of  the  width  of  horny  beak. 

f.  Tadpole  large  (65  mm  ),  transforming  at  13  -2.4  mm.;  teeth  4/4,  rarely 
5/4  or  3/4,  upper  fringe  of  papillae  broken  in  middle  by  a  row  of  teeth; 
eye  1.1-1.3  nearer  tip  of  snout  than  to  the  spiracle,  av.  i  i,  muscular 
part  of  tail  1.15- i  66  in  depth  of  tail,  av.  1.43 ,  width  of  body  in  its  own 
length  1.55  -1.9,  av.  1.66,  depth  of  body  1.05-1.15  in  body  width,  av. 
1.09,  spiracle  j  15  -i  6  nearer  eye  than  base  of  hind  legs  or  vent,  av. 
1.37,  internasal  space  1.5-175  in  interorbital  space,  av.  1.6,  third 
lower  labial  row  broken  in  middle,  median  interval  of  second  lower 
row  broad,  last  row  of  teeth  1.75-10  times  in  next  to  last  row  of 
teeth  and  usually  i  .7-1.0  times  in  mandibles,  second  upper  row  usually 
broken  in  middle.  Egg  mass  a  loose  cylinder,  many  eggs  on  stalks. 
Montana  to  Texas  and  westward  to  Pacific  Coast  States. 

Scaphtopus  hammondii 

(g.  Upper  labial  mandible  with  beak,  lower  labial  mandible  with 
notch.  Scaphtopus  h.  hammondii 

Plate  XVlII-3,5 

gg  Upper  labial  mandible  without  beak,  lower  labial  mandible  with- 
out notch.  Scaphtopus  h.  bombifront 
After  Bragg) 

[ggg.  Probably  belongs  with  gg.  Scaphtopus  h.  tnttrmontanut] 

ff.  Mature  tadpole  small  (2.1-2.7  mm.),  transforming  at  7  5-11.5  mm., 
upper  fringe  of  papillae  is  conspicuously  broken  in  middle  by  a  row 
of  teeth  or  not  so  broken  and  the  first  row  absent,  last  lower  row  of 
teeth  1.1-1  4  in  width  of  mandibles. 

g.  Teeth  usually  4/5,  sometimes  3/4,  interval  between  lateral  parts 
of  third  upper  row  i  5  -3  o  in  either  lateral  row  or  5.0  in  width 
of  mandibles,  interval  between  lateral  portions  of  fourth  row  5  o 
times  in  width  of  mandibles,  second  upper  row  normally  broken. 
Eggs  tandcmhke,  or  Bufo-likc  or  lattice  work.  (Compiled  after 
Bragg 0  Oklahoma  and  Arkansas  to  Texas.  Scaphiopus  h.  hurteni 

gg.  Teeth  usually  4/4,  rarely  5/4,  3/4,  or  possibly  1/4,  interval  be- 
tween lateral  parts  of  third  upper  row  equal  to  or  1/3  either  lateral 
portion  of  this  row  or  1.5  times  in  mandible,  interval  between 
lateral  portions  of  fourth  upper  row  1.3  in  width  of  mandibles; 
second  upper  row  normally  unbroken,  eye  i. 0-1.6  nearer  tip  of 


GENERAL  ACCOUNT 


57 


snout  than  to  the  spiracle,  average  1.3,  muscular  part  of  tail 
1.1-3.33  'm  depth  of  tail,  av.  1.7;  width  of  body  in  its  own 
length  1.3-1.6,  av.  1.4,  depth  of  body  1.15-1.55  in  body  width, 
av.  1.35;  spiracle  1.5-1.1  nearer  eye  than  base  of  hind  legs  or 
vent,  av.  1.85,  mternasal  space  1.45-1.3  in  mterorbital  space, 
av.  i.o,  third  lower  labial  row  continuous;  median  interval  of 
second  lower  row  very  narrow.  Last  lower  row  1.5-3.0 
times  in  next  to  lower  row  of  teeth;  second  upper  row  not  or 
barely  broken  in  middle,  egg  mass  an  irregular  cylinder,  at  first 
bandlike.  Texas  to  California  and  Mexico.  Scaphtopus  coucbii 

dd.  Labial  teeth  1/3;  papillae  confined  to  the  sides  of  the  labium  (or  lower 
half  only  in  Bufo  punctatus),  upper  and  lower  edges  toothed,  papillar 
border  on  each  side  emarginate,  eyes  slightly  nearer  lateral  outline  than 
middorsal  line,  above  lateral  axis,  spiracle  small,  a  porehke  opening, 
e.  Tadpoles  14-18  mm.  in  length  (a  few  forms  35-56  mm  ) 

BUFONIDAE 

f.  Papillae  only  on  lower  half  of  lateral  margin,  or  a  slight  marginal 
row  of  4  to  6  papillae  on  upper  half,  no  inner  papillae  normally, 
third  lower  row  of  teeth  equal  to  the  first  row  of  lower  labial 
teeth,  median  space  between  lateral  parts  of  the  second  row  of 
upper  labial  teeth  1-3  times  in  either  lateral  row,  horny  beak 
1.1-1.5  in  third  lower  labial  row,  tadpoles  to  15  mm  One  of  the 
blackest  Bufo  tadpoles,  tail  musculature  evenly  dotted  with  black, 
venter  with  light  grayish  vmaceous  spots,  eggs  single  or  film  or 
scattered  mass  on  bottom,  not  in  files.  Central  Texas  to  California, 
Utah,  and  Lower  California.  Bufo  punctatus 

Plate  XIII  5 

ff.  Papillae  on  upper  and  lower  halves  of  lateral  labial  margin,  some 
inner  papillae. 

g.  Bicolor  tadpoles  distinctly  lighter  below,  not  black  tadpoles, 
upper  tail  fin  highly  arched,  horny  beak  1.3-1.5  in  first  or 
second  lower  row  of  teeth. 

h.  Light  grayish  olive,  dark  olive-buff  or  clay  color,  the 
lightest  of  our  toad  tadpoles,  in  life  intestine  does  not  show 
through  the  skin  of  the  belly  (shows  in  preserved  speci- 
mens), third  lower  row  of  labial  teeth  short,  1.8-1.4  in 
first  lower  row,  third  lower  row  1.5-1  o  times  in  the  horny 
beak  or  1.6-1.8  m  longest  lower  row,  median  space  between 
lateral  parts  of  the  second  row  of  upper  labial  teeth  large, 
1. 1-1.0  times  greater  than  either  lateral  part,  depth  of  tail 
in  length  of  tail  3.0-3.5;  horny  beak  1.3-1.4  in  first  or 


58  HANDBOOK  OF  FROGS  AND  TOADS 

second  row  of  teeth;  lower  loop  of  papillae  to  end  of  third  lower  but 
not  under  it,  usually  not  two  rows;  tail  musculature  with  dark 
vinaceous  drab  band  to  tail  tip;  below  this  a  pale  vinaceous  pink 
band,  belly  pale  vinaceous  pink;  eggs  in  files  or  strings,  brown  or 
buffy  brown,  cream  cr  straw  yellow  below;  no  inner  tube.  Texas  to 
Mexico,  Arizona.  Bufo  compactilis 

Plate  XIII-6 

(hh.  Doubtless  Bragg's  (1936)  Bufo  cognatus  belongs  in  g.  Dorsal  surface 
mottled  brown  and  gray  silvery  areas  interwoven  with  black  ones. 
The  ventral  surface  markedly  lighter  than  the  dorsal.  The  viscera 
(in  mature  tsdfcles)  cnly  slightly  or  not  at  all  visible,  lower  tail  fin, 
clear  or  almost  so,  third  lower  rcw  of  teeth  1.5-1  8  times  in  horny 
beak,  or  1.5-3.0  in  longest  lower  row,  median  space  in  second  upper 
row  (none  says  Bragg,  1.0-1.2.  in  either  lateral  part  says  Smith), 
depth  of  tail  in  length  of  tail  1.5,  eggs  in  single  row  sometimes 
double,  an  inner  gelatinous  envelope  and  an  outer  tougher  envelope 
or  tube,  somewhat  scalloped  in  outward  appearance.  (After  A.  N. 
Bragg,  Okla.,  1936,  p.  19.)  Minnesota  to  Texas  and  southern  Cali- 
fornia. Bufo  cognatus) 

gg.  Elack  or  blackish  tadpoles,  in  life  the  intestine  shows  through  the  skin 

of  the  belly,  third  lower  rcw  of  labial  teeth  long,  1.1-1.6  in  first  lower 

row,  third  lower  row  1.0-1.5  in  horny  beak;  median  space  between 

lateral  parts  of  the  second  row  of  upper  labial  teeth  small,  contained 

1.0-4.9  times  in  either  lateral  half,  not  greater  than  lateral  half. 

h.  Papillae  very  faint,  minute,  at  times  hard  to  see  but  present;  third 

lower  rcw  of  labial  teeth  not  equal  to  first  lower  rcw  of  teeth,  but 

equal  to  or  greater  than  horny  beak,  horny  beak  1.1-1.5  in  upper 

fringe,  1.3-1.5  in  first  lower  row;  median  space  of  second  upper  row 

1-4  times  in  either  lateral  part  of  it;  upper  edge  of  tail  musculature  with 

8- 10  black  bars  with  intervening  pale  olive-buff  areas;  irregular  black  or 

brown  band  on  tail  with  cartridge  buff  or  tilleul  buff  below  it;  tadpoles  to 

2^  mm.  Eggs  in  files,  inner  tube  present,  one  or  two  rows  of  eggs. 

Louisiana  to  Costa  Rica.  Bufo  valliceps 

Plate  XIII-7 

hh.  Papillae  plainly  visible. 

i.  Tadpoles  of  Rockies  to  eastward.  Tadpoles  usually  less  than  30 

mm.,  third  lower  row  of  labial  teeth  1.1-1.5  *n  fifst  row- 

j.  Tadpole  to  14  mm.;  horny  beak  in  upper  fringe  1.75,  in  first  or 

second  lower  row  1.5 ,  horny  beak  equal  to  or  less  than  the  third 

lower  row  of  teeth;  median  space  between  two  parts  of  second 


GENERAL  ACCOUNT 


59 


upper  row  1.4-1.1  in  cither  lateral  part;  third  lower  row  of  teeth  in  first 
lower  row  i  .1-1.4; two  or  more  rows  of  strong  papillae  from  end  of  upper 
fringe  to  end  of  third  lower  row,  sometimes  3-5  rows  at  side  of  labium; 
lower  loop  of  papillae  far  below  level  of  third  row  and  with  at  least  two 
rows  of  papillae,  mouth  in  interorbital  space  1.0-1.5,  av-  I-I7>  mouth 
larger  than  internasal  space  1.1-1.6,  av.  1.36;  depth  of  tail  in  tail  length 
1.4-4.5,  av.  3.17,  eye  nearer  snout  than  spiracle  1.0-1.57,  av.  1.1;  nostril 
nearer  eye  than  snout  1.1-1.8,  av.  1.48.  Egg  mass  long  file,  inner  tube 
with  no  partitions.  North  Carolina  to  Florida  to  Louisiana. 

Bufo  temstris 
Plate  XII -ii 

jj.  Tadpole  to  17  mm.;  horny  beak  in  upper  fringe  1.1-1.5,  *n  ^rst  or  second 
lower  row  of  teeth  1.1-1.1  or  1.1-1.3;  horny  beak  greater  than  third  row 
of  lower  labial  teeth;  median  space  1.15-1.0,  1.0-4  °>  or  I-3~3-°  *n  either 
lateral  part,  third  lower  row  of  teeth  1.3-1.5  or  1.4-1.5  in  first  lower 
row;  one  row  of  weak  papillae  from  upper  fringe  to  end  of  third  lower 
row  of  teeth  with  a  few  scattering  papillae  at  the  side  of  the  labium, 
lower  loop  with  only  two  or  three  scattering  papillae  beside  the  outer 
row  of  weak  papillae. 

k.  Mouth  in  interorbital  distance  0.77-1.0,  av.  0.91;  horny  beak  in  upper 
fringe  1.1-1.4,  horny  beak  in  first  or  second  row  1.1-1.1  times;  third 
row  in  first  lower  row  1.3-1.5,  depth  of  tail  in  tail  length  1.15-1.7, 
av.  1.97,  spiracle  nearer  eye  than  vent  1.04-1.54,  av.  i  18;  eye  nearer 
snout  than  spiracle  i  0-1.17,  av.  1.16,  mouth  larger  than  internasal 
space  1.4-1 i,  av.  i  76;  papillae  of  lower  labial  loop  do  not  extend 
under  the  end  of  the  third  row  of  labial  teeth,  tail  musculature  in  tail 
depth  1.16-1.66,  av.  1.04,  internasal  space  1.1-1.8  in  interorbital  dis- 
tance, av.  1.6,  spiracle  1.05-1.55  nearer  eye  than  vent,  av.  1.18.  Egg 
mass  long  file,  partitions,  inner  tube  present.  Eastern  North  America 
from  Hudson  Bay  southeast.  Bufo  a.  amcrtcanus 

Plate  XII-io 

kk.  Mouth  in  interorbital  distance  0.9-1.5,  horny  beak  1.4-1.9  in  first 
upper  fringe  of  teeth,  horny  beak  in  first  or  second  row  1.1-1.4; 
depth  of  tail  in  tail  length  1.75-3.8. 

1.  Mouth  in  interorbital  distance  1.07-1.5,  horny  beak  in  upper  fringe 
1.4-1.5,  horny  beak  in  first  or  second  row  1.1-1.3,  third  lower  row 
1.3-1.6  in  first  lower  row;  depth  of  tail  in  tail  length  1.88-3.83, 
av.  3.33,  spiracle  nearer  eye  than  vent  1.15-1.7,  av.  1.45,  eye  nearer 
snout  than  spiracle  1.0-1.6,  av.  1.16;  mouth  larger  than  internasal 
space  i .  i-i . 83 ,  av.  i  .47,  papillae  of  lower  labial  loop  extend  slightly 
or  not  at  all  under  the  end  of  the  third  lower  row,  tail  musculature 
in  tail  depth  1.6-1.3,  av.  1.85;  internasal  space  1.5-1.16  in  inter- 


60  HANDBOOK  OF  FROGS  AND  TOADS 

orbital  space,  av.  1.86;  spiracle  1.15-1.7  nearer  eye  than  vent  or 
base  of  hind  legs,  av.  1.45.  Egg  mass  long  file,  no  inner  tube, 
sometimes  two  rows  of  eggs.  Massachusetts  to  Missouri  on  the 
north  and  South  Carolina  to  Texas  on  the  south.  Bufo  w.  fowleri 

Plate  XII-7 

11.  Horny  beak  in  upper  fringe  1.8-1.9;  in  first  or  second  lower  row 
of  teeth  1.6;  third  lower  row  1.3-1.4  in  first  lower  row,  depth 
of  tail  in  length  of  tail  1.75;  spiracle  nearer  eye  than  vent  1.15; 
eye  nearer  snout  than  spiracle  i.o-i.i  or  equidistant,  papillae  of 
lower  labial  loop  slightly  extends  under  the  ends  of  the  third 
lower  row;  tail  musculature  in  tail  depth  1.1-1.5;  egg  mass  a 
long  file,  no  inner  tube,  sometimes  two  rows  of  eggs.  (After  K. 
A.  Youngstrom  and  H.  M.  Smith,  Kan.,  1936,  p.  633.)  Western 
Iowa  south  through  Texas  into  Mexico  and  west  to  Idaho  and 
Imperial  Valley,  California.  Bufo  w.  woodbousii 

ii.  Tadpoles  of  Pacific  Coast  area.  Tadpoles  often  over  30  mm.  (maximum 
17-56  mm  ,  2.7  canorus,  33  exsul,  19  nelson^  44  borcas,  56  b.  halophtlus — 
boreas  group).  Third  lower  row  of  labial  teeth  1.4  or  1.5-1.0  in  first  lower 
row. 

j.  Smaller  tadpoles  (17  mm.  canorus,  35  mm.  exsuf).  Median  space  in 
second  upper  row  of  teeth  almost  equal  to  or  1.1  times  in  either  lateral 
portion  of  this  row,  third  lower  row  1.5-1.0  in  first  and  second  lower 
rows,  1.1-1.5  *n  first  upper  fringe. 

k.  Third  lower  row  1.5  in  first  lower  row,  i.o  in  second  lower  row  or 
1.1-1.3  ifl  upper  fringe.  (After  Storer.)  Yosetnite  region. 

Bufo  canorus 

kk.  Third  lower  row  1.4  in  first  or  second  lower  row  or  equal  to  or 
1.5  in  first  upper  fringe.  Deep  Springs,  Calif.  Bufo  exsul 

}].  Larger  tadpoles  (19  mm.  nelsoni,  55  mm.  b.  halophtlus}.  Median  space 
in  second  upper  row  of  teeth  1.0-3.7  in  either  lateral  rows,  third  lower 
row  1.4-1.0  in  first  or  second  lower  row. 

k.  Median  space  in  second  upper  row  1.0^-1.5  times  in  either  lateral 
portion  of  this  row  or  1.5-3.0  in  horny  beak;  third  lower  row  of 
teeth  i  times  in  first  or  second  lower  row  of  teeth.  Southern  Nevada 
into  California.  Bufo  b.  nelsoni 

kk.  Median  space  in  second  upper  row  of  teeth  3-2.-3.7  in  either  lateral 
row  or  3.8  in  horny  beak;  third  lower  row  1.4  in  first  or  second 
lower  row  or  1.5-1.6  in  first  upper  fringe.  (After  T.  I,  Storer.) 


GENERAL  ACCOUNT  61 

Montana  and  Colorado  to  northern  California 
and  British  Columbia,  possibly  Alaska. 

Bufo  b.  halophtlus 

[The  characters  used  for  this  boreas  group  are 
trivial  and  not  to  be  given  too  much  significance.] 

ee.  Tadpoles  55  or  56  mm.  (body  2.3  and  tail  33;  see  Storer  for  descrip- 
tion). Bufo  b.  balophtlus 

Anus  dextral,  spiracle  distinctly  lateral  on  or  near  body  axis. 

d.  Papillary  border  on  side  with  an  emargination,  tadpoles  50-149  mm. 
in  length;  papillary  fringe  on  upper  labium  extends  not  at  all  inward 
beyond  the  end  of  the  upper  fringe  of  teeth  or  only  1/8  to  1/16  of  the 
length  of  the  fringe;  length  of  horny  beak  in  upper  fringe  of  teeth 
i.o^-i  5  times;  labial  teeth  1/3  or  3/3  or  3/4,  rarely  to  7/6. 

RANIDAE 

e.  Labial  teeth  7/6  to  3/3,  not  customarily  1/3,  tadpoles  47-88  mm. 

f.  Labial  teeth  7/6  or  6/6,  depth  of  tail  in  length  often  3.3-4.1. 

(See  T.  I.  Storer  for  description  )  Coastal  California  and  Oregon. 

Rana  b.  boy  It  i 
ff.  Labial  teeth  1/4,  3/4,  4/3,  or  5/3,  not  1/3. 

[R.  aurora  aurora,  labial  teeth  3/4,  belongs  here;  no  detailed  de- 
scription on  record,  probably  some  R.  a.  draytom  also  fall  here 
though  some  are  1/3  (T.  I.  Storer).] 

g.  Labial  teeth  5/3  or  4/3,  tadpoles  large  to  88  mm.,  tadpole 
tail  very  prominently  spotted  and  mottled,  all  three  lower 
labial  rows  long,  median  space  of  second  upper  row  1.3-1.5 
in  either  lateral  section  of  this  row,  tail  tip  pointed  or 
rounded,  depth  of  tail  in  tail  length  3.3-3.4  times,  spiracle 
1.1-1.3  ne&rer  vent  than  tip  of  snout,  nostril  1.3-1.8  nearer 
eye  than  snout,  spiracle  1.8-1.0  nearer  eye  than  vent.  New 
Mexico,  southern  Arizona  to  central  Mexico. 

Rana  tarabumarat 
Plate  CXXII-i,6 

gg.  Labial  teeth  3/4,  tadpoles  49-71  mm.  in  length.  Dorsal  crest 

very  high  extending  to  vertical  of  the  spiracle  (Rana  sylvattca) 

or  tail  broader  nearer  its  tip  than  at  its  body  insertion, 

rounded,  spatulate,  or  elliptical  (R.  b.  surrai);  upper  fringe 

of  teeth  about  1.5  times  the  length  of  the  horny  beak. 

h.  Tadpole  to  49  mm.;  dorsal  crest  very  high,  extends  on  the 

body  to  vertical  of  the  spiracle;  tail  tip  acuminate,  tail 

musculature  begins  to  taper  at  once;  dorsal  crest  higher 

in  cephalic  half;  depth  of  tail  in  length  of  tail  1.9-3.1, 

av.  1.5;  spiracle  1.5-1.0  nearer  base  of  hind  legs  or  vent 


62  HANDBOOK  OF  PROGS  AND  TOADS 

than  tip  of  snout,  av.  1.75;  nostril  equidistant  from  snout  and 
eye;  internasal  space  in  interorbital  space  1.7-1.9,  av.  1.5;  depth 
of  body  1.08-1.13  in  body  width,  av.  1.14;  depth  of  body  1.66- 
z.o  in  body  length,  av.  1.78,  second  upper  lateral  row  about 
1/5  of  the  upper  fringe  of  teeth,  the  median  space  between  the 
lateral  portions  of  the  second  row  0.4-0.8  times  either  lateral 
portion;  the  third  upper  row  is  1/4  to  1/9  of  the  upper  fringe; 
fourth  lower  row  of  teeth  is  6/n  to  7/11  of  the  first  lower  row; 
spiracle  1.0-1.47  of  the  first  lower  row;  spiracle  1.0-1.48  nearer 
eye  (5.0-6.0  mm.)  than  base  of  hind  legs  or  vent  (4  5-7.6  mm.). 
Egg  mass  submerged,  globular.  Ontario  to  Nova  Scotia,  south 
to  South  Carolina,  west  to  South  Dakota  and  south  to  Arkansas. 

Rana  s.  sylvatica 

hh.  Tadpole  to  71  mm.;  dorsal  crest  low  extending  on  body  to  a 
vertical  twice  nearer  hind  legs  than  spiracle;  tail  tip  rounded, 
elliptical,  or  spatulate;  tail  musculature  for  an  inch  or  more 
does  not  taper;  dorsal  crest  narrow  in  cephalic  half,  depth  of 
tail  in  length  of  tail  1.5-5.7,  av.  4.0;  depth  of  body  1.1-1.8  in 
body  width,  av.  1.43,  depth  of  body  in  body  length  1.0-3.0, 
av.  1.315,  spiracle  1.0-1.33  nearer  base  of  hind  legs  or  vent  than 
tip  of  snout,  av.  1.17;  spiracle  1.45-1.375  nearer  eye  (4.0-8.0 
mm.)  than  base  of  hind  legs  or  vent  (5  8-n.o),  av.  1.05;  young 
tadpoles  occasionally  with  teeth  1/4,  rarely  3/3;  nostril  i.o- 
1.75  nearer  the  eye  than  snout,  internasal  space  1.3-1.0  in  inter- 
orbital  space,  av.  1.5;  second  upper  lateral  row  1/4  to  1/9  of 
the  upper  fringe;  median  space  between  lateral  portions  of  sec- 
ond upper  row  1.0-1.3  times  either  lateral  part;  the  third  upper 
row  is  1/9  to  i/n  of  the  upper  fringe;  the  fourth  lower  row  of 
teeth  is  1/3  to  1/4  of  the  first  lower  row.  The  Sierras,  California. 

Rana  b.  sicrrac 

ggg-  Labial  teeth  1/4;  fourth  lower  labial  row  1/4  of  length  of  other 
three;  median  space  in  second  row  small.  (After  J.  R.  Slater,  Wash.) 
Tadpoles  40-50  mm.  Northern  Oregon  into  Central  Washington. 

Rana  cascadae 
ee.  Labial  teeth  1/3,  occasionally  3/3,  rarely  1/3. 

(We  have  seen  no  tadpoles  of  R.  prcttosa  or  R.  p.  lutcwentns  in  the  field; 
we  have  seen  preserved  tadpoles  with  teeth  3/3.  "In  prcttosa  there  are 
four  rows  of  labial  teeth,  one  very  short  upper  row  and  three  lower  rows. 
In  luttiventns  there  are  five  rows,  two  long  upper  rows  and  three  lower, 
as  figured  by  Thompson  (1913).  The  first  lower  row  in  lufetwnfris  is  longer 
than  the  analogous  one  in  pretiosa"  (A.  Svihla,  Wash.,  1935,  p.  i").) 
Tadpoles  74-150  mm.  in  length;  dorsal  crest  not  extending  to  vertical  of 
the  spiracle  but  usually  just  ahead  of  the  buds  of  the  hind  legs;  tail 
always  elliptical  not  spatulate;  upper  fringe  of  teeth  equal  to  or  slightly 
larger  than  (never  1.5  times)  the  horny  beak. 


GENERAL  ACCOUNT  63 

f.  Tadpoles  74-84  mm.;  tadpoles  usually  transform  the  same  season  they  are 
born;  transformation  sizes  18-30  mm.,  av.  2.4  mm.  (except  R.  arcolata 
group),  tadpoles  (except  in  R.  capita  or  arcolata  group)  not  strongly  pig- 
mented  on  belly,  viscera  plainly  or  slightly  showing  through  skin  (in 
spirit  specimens).  Egg  mass  globular  or  plinthlike  beneath  surface  of 
water. 

g.  Belly  strongly  pigmented  or  somewhat  pigmented,  in  spirits  looking 
white  and  viscera  not  visible  or  but  slightly  visible;  tail  covered  with 
large  prominent  dark  spots;  transformation  2.5-35  mm- 
h.  Nostril  equidistant  eye  and  snout,  av.  1.15 ;  spiracle  equidistant  vent 
and  snout,  nearer  eye  (1.5)  than  vent,  intestines  show  through  belly 
but  not  quite  like  R.  piptens;  median  space  between  second  upper 
row  of  teeth  1.15-1.5  times  in  length  of  cither  lateral  part;  third 
lower  row  0.40-0.66  of  the  second  lower  row;  spotting  not  so  promi- 
nent as  in  R.  capto  but  heavier  than  in  R.  pipens  group;  inadequate 
characterization.  Southwestern  Ohio  to  Kansas  and  Oklahoma  and 
northeastern  Louisiana.  Rana  arcolata  circulosa 

hh.  Nostril  1.0-1.5  flearer  eye  than  snout,  av.  1.15;  eye  1.0-1.3  nearer 
spiracle  than  snout,  av.  1.11,  median  space  between  the  second  up- 
per row  of  labial  row  1.0-1.0  times  the  length  of  either  lateral  part 
of  this  row,  third  lower  row  0.33-0.66  shorter  than  the  first  or 
second  rows,  tail  covered  with  large  prominent  dark  spots;  belly 
strongly  pigmented  (in  spirits  it  looks  white),  viscera  not  visible; 
depth  of  tail  in  length  of  tail  i.fr  3.5,  av.  3.0.  North  Carolina  to 
Florida.  Rana  capita 

Plates  XI-i,i;  XV~}-8 

gg.  Belly  not  strongly  pigmented,  in  spirits  dark,  viscera  shows  through 
skin,  tail  in  R.  piptens  group  with  fine  flecks  or  small  spots,  or  in  R. 
palustris  heavily  washed  with  purplish  or  purplish  black,  transforma- 
tion sizes  averaging  smaller,  18-31  or  31  mm.  [Rana  aurora  Jraytonti. 
Body  1. 1-1.1  in  length  of  tail;  depth  of  tail  in  length  of  tail  1.5-3.1  in 
its  length.  (See  T.  I.  Storer's  description.)] 

h.  Median  space  in  second  upper  labial  row  1-4  times  the  length  of 
either  lateral  part;  third  row  of  lower  labial  teeth  0.33-0.66  shorter 
than  the  first  or  second  rows,  usually  at  least  0.50,  eye  nearer  the 
snout  than  spiracle  or  equidistant,  nostril  nearer  eye  than  tip  of 
snout;  depth  of  tail  in  length  of  tail  1.3-3.1,  av.  1.7;  spiracle  1.5-1.8 
nearer  eye  than  snout,  av.  1.63.  Hudson  Bay  to  Texas  and  eastern 
states.  Rana  palustris 

Plate  XI-I4 


64  HANDBOOK  OF  FROGS  AND  TOADS 

hh.  Median  space  0.5-1.0  or  1.0-1.5  times  either  lateral  part  of  the 
second  upper  row,  third  row  of  labial  teeth  o  2.2.  or  0.185-0.33 
shorter  than  the  first  lower  row,  depth  of  tail  in  length  of  tail 
1.0-1.8  or  1.7-3.4. 

i.  Median  space  of  second  upper  row  0.5-1.0  times  either  lateral 
part,  third  lower  row  of  teeth  0.185-0  33  shorter  than  the  first 
lower  row,  nostril  to  snout  equal  to  nostril  to  eye,  eye  1.15-1.3 
nearer  tip  of  snout  than  spiracle,  av.  1.1;  body  length  in  tail 
length  1.35-1.66,  av.  1.5,  mouth  0.9-1.6  larger  than  internasal 
space,  depth  of  tail  in  length  of  tail  1.0-1.88,  av.  1.65,  tail  crest 
usually  with  wide  prominent  dark  spots,  greatest  length  of  tad- 
pole 74  mm  ;  spiracle  1.1-1.86  nearer  eye  than  snout,  av.  1.46. 
Southeastern  states  to  Texas,  Oklahoma,  and  Indiana. 

Rana  piptms  sphcnoccphala 
Plate  XI-9,io 

ii  Median  space  1.0-1.5  times  either  lateral  part,  third  lower  row 
o  11  shorter  than  the  first  lower  row,  nostril  i.i-i  5  nearer  the 
eye  than  snout,  eye  nearer  spiracle  than  snout  1.1-1.3;  body 
length  in  tail  length  1.3-1.0,  av.  1.7,  mouth  larger  than  inter- 
nasal  space,  depth  of  tail  in  length  of  tail  1.7-3  4>  av-  3-°>  tail 
crest  usually  translucent  with  fine  spots  or  pencilmgs,  greatest 
length  of  tadpole  84  mm.,  spiracle  i  4-1.86  nearer  eye  than 
snout,  av.  1.59.  North  America  east  of  Sierra  Nevada  and  south- 
ward into  Mexico.  Rana  p.  ptptent 

Plate  XI-n 

ff.  Tadpoles  83-141  mm  ,  tadpoles  usually  winter  over  at  least  one  season; 

transformation  sizes  18  59  mm.  (except  R.  vtrganpes  15-35  mm  >  possibly 

R.  Jishcri),  tadpoles  usually  with  strongly  pigmented  bellies,  viscera  not 

plainly  showing  through  the  skin  (in  spirit  specimens). 

g.  Tadpole  with  prominent  continuous  black  crest  margins  and  a  black 

musculature  band,  belly  bluish;  tadpoles  to  95  mm  ,  young  tadpoles 

black  with  transverse  yellowish  band  on  the  body,  spiracle  0.86-1.1 

nearer  vent  or  base  of  hind  legs  than  snout,  average  i.o,  spiracle  0.85- 

1.1  nearer  eye  than  base  of  hind  legs  or  vent,  i.e.,  usually  equidistant, 

eye  equidistant  from  spiracle  and  tip  of  snout,  muciferous  crypts  very 

distinct,  spiracle  below  lateral  axis;  tail  tip  acuminate,  second  upper 

labial  row  in  upper  fringe  1/3  to  1/4,  upper  fringe  distinctly  greater 

than  horny  beak,  median  space  between  two  parts  of  second  upper 

labial  row  i  to  i\%  of  either  lateral  part,  third  lower  labial  row  equal 

to  horny  beak,  third  lower  labial  row  longer  than  single  row  of  lower 

papillae,  third  lower  labial  row  1/4  to  1/5  shorter  than  first  lower  row. 

Eggs  not  described.  South  Carolina  to  Florida  and  Louisiana. 

Rana  beckscheri 
Plates  XI-7,8;  XIV-i,i;  XCVIII-i,6 


GENERAL  ACCOUNT  65 

gg.  Tadpole  without  black  crest  margins  or  lateral  band,  belly  white,  car- 
tridge buff  or  yellow  to  maize  yellow,  no  transverse  yellow  band  in 
young  tadpoles,  spiracle  nearer  vent  than  snout  1.1-1.8;  spiracle  to  eye 
rarely  less  than  1.2.5  greater  than  spiracle  to  vent,  eye  nearer  tip  of  snout 
than  spiracle  1.0-1.4,  second  upper  labial  row  to  upper  fringe  1/4  to  1/15, 
upper  fringe  equal  to  or  slightly  greater  than  horny  beak,  median  space 
in  either  lateral  part  ij^  to  u;  third  lower  labial  row  much  less  than 
horny  beak,  third  lower  row  much  shorter  than  or  equal  to  single  row 
of  lower  labial  papillae. 

h.  Tadpoles  to  140  mm.,  eye  well  above  lateral  axis,  muciferous  crypts 
indistinct;  spiracle  just  below  lateral  axis,  spiracle  1.08-1.44  nearer 
base  of  hind  legs  or  vent  than  tip  of  snout,  av.  1.2.6;  depth  of  tail  in 
length  of  tail  2-4-3. 5»  av*  x-8>  ta^  C1P  obtuse;  second  upper  row  in 
upper  fringe  1/4  to  1/7,  median  space  in  second  upper  row  \Yi  in 
either  lateral  part,  third  lower  row  in  first  lower  1/4  to  1/5  shorter; 
teeth  1/3,  rarely  3/3.  Transformation  size  43-59  mm.  Egg  mass, 
surface  film.  North  America  east  of  Rockies.  Rana  catesbciana 

Plate  XCIV-3 

hh.  Tadpoles  to  83-110  mm.;  eye  on  or  just  above  lateral  axis,  tail  tip 
acute  or  acuminate  (rounded  in  R.  fishcrt)\  teeth,  1/3,  second  upper 
row  fringe  1/6  to  1/15,  median  space  in  second  upper  row  1.5-11  in 
either  lateral  part,  third  lower  row  in  first  lower  row  1/2.  to  1/3 
shorter. 

i.  Depth  of  tail  in  length  of  tail  1.45-1.8,  average  1.7;  tail  tip  acumi- 
nate, dorsal  crest  equal  to  or  less  than  tail  musculature,  muciferous 
crypts  indistinct,  spiracle  1.08-1.44  nearer  vent  than  snout;  mouth 
in  interorbital  distance  1.5-2.. 37,  av.  1.94,  internasal  space  in  in- 
terorbital  distance  1.8-1.6,  av.  1.16;  second  upper  row  1/6  to  1/8 
of  the  upper  row,  median  space  of  the  second  upper  row  2.%  to 
4!^  times  either  lateral  row,  third  lower  row  i  5  less  than  horny 
beak,  much  shorter  than  single  row  of  lower  labial  papillae  and 
1/2.  shorter  than  first  lower  row  of  teeth,  first  row  of  lower  teeth 
equal  to  horny  beak.  Transformation  size  31  or  37  to  48  mm.;  egg 
mass,  a  surface  film.  Georgia  to  Florida  to  Louisiana.  Rana  grylto 

Plates  XI-5,6,  XIV -5 

ii.  Depth  of  tail  in  length  of  tail  1.5-4.7,  avs.  3.1-3.87,  tail  tip  acute, 
dorsal  crest  less  than  tail  musculature,  muciferous  crypts  distinct, 
species  1.07-1.8  nearer  vent  than  snout,  mouth  in  interorbital  dis- 
tance 1.3-1.8,  av.  1.5,  internasal  space  in  interorbital  space  i  15- 
i.o,  avs.  1.6-1.75;  second  upper  row  1/6  to  1/15  of  the  first  upper 
row;  first  lower  row  of  teeth  equal  to  or  greater  than  horny  beak, 
spiracle  1.1-1.8  nearer  vent  than  snout. 


66  HANDBOOK  OF  FROGS  AND  TOADS 

j.  Tadpoles  to  99  mm.,  transformation  size  to  19-38  mm.;  depth  of  tail  in 
length  of  tail  3.1^4.7,  av.  3.87;  spiracle  just  touches  lateral  axis;  eye  just 
above  lateral  axis,  spiracle  1.06-1.38  nearer  eye  than  base  of  hind  legs 
or  vent,  av.  1.2.4,  spiracle  1.15-1.6  nearer  eye  than  vent,  1.45,  mouth  in 
interorbital  distance  1.3-1.75,  av.  1.55,  width  of  body  in  its  own  length 
1.3-1.1,  av.  1.55,  third  lower  labial  row  of  teeth  1.15  less  than  horny 
beak,  about  equal  to  single  row  of  lower  labial  papillae,  1/1  shorter  than 
first  lower  row,  sometimes  a  row  of  inner  papillae  below  the  third  lower 
row  of  teeth,  median  space  in  second  upper  labial  row  3.5-4.5  times  either 
lateral  portion;  second  upper  row  1/6  to  1/15  of  the  upper  fringe,  belly 
straw  yellow,  colonial  buff,  or  deep  colonial  buff,  tail  with  round  car- 
tridge buff  or  pinkish  cinnamon  spots,  no  black  line  in  dorsal  crest  as  in 
R.  grylio  or  R.  virgattpcs.  Eggs  in  a  compact  submerged  mass.  Hudson 
Bay  to  Minnesota,  New  York  to  New  England.  Rana  scptcntrionalts 

Plates  XI-n;  XIV-3 

j).  Tadpoles  to  91  mm  ,  transformation  sizes  at  15-38  mm.,  depth  of  tail  in 
length  of  tail  1.5-3  7>  spiracle  just  below  lateral  axis,  eye  on  lateral  axis, 
third  lower  labial  row  of  teeth  1.5-  1.2.5  ^ess  than  horny  beak,  much 
shorter  than  single  row  of  lower  labial  papillae,  almost  1/2.  shorter  than 
first  lower  row,  median  space  of  second  upper  row  6-n  times  the  length 
of  either  lateral  portion;  second  upper  row  1/15  to  1/15  of  the  first  upper 
row. 

k.  Spiracle  nearer  vent  than  snout  1.35-1  8,  mouth  1.3-1.8  times  in  inter- 
orb'tal  distance,  av.  i  5,  width  of  body  in  its  own  length  1.15-1.7, 
av.  1.47;  belly  deep  cream  color,  tail  green,  mottled  with  brown  and 
covered  with  line  yellow  spots.  Egg  mass,  surface  film.  Canada  to 
Louisiana  to  Florida  to  New  England.  Rana  damttans 

Plates  XI-3,4,  XIV-7 

kk.  Spiracle  nearer  vent  than  snout  1.07-1.45,  av.  1.13-1.3,  mouth  i.o- 

1.37  in  interorbital  distance,  av.  1.11. 

1.  Teeth  1/3  or  1/3 ,  no  inner  papillae  or  few  inner  papillae  from  end  of 
the  upper  row  to  the  end  of  the  lower  labial  row,  no  row  of  finer 
papillae  below  third  lower  row  of  teeth,  spiracle  nearer  eye  than 
vent  1.1-1.75,  'lv-  *  43'  nostril  nearer  eye  than  snout  1.1-1.9,  av- 
1.5,  tadpoles  medium  ^our  material,  41  mm.,  Stanford  University 
material,  83  mm.),  width  of  body  in  body  length  1.6-1.0,  av.  1.77; 
second  upper  row  1/6  to  i  /8  of  the  first  upper  row,  belly  pure  white 
or  pale  cinnamon  pink,  tail  musculature  with  black  clusters  out- 
lining cartridge  buff  areas,  upper  tail  crest  sometimes  reticulated 
with  black  dots.  Lower  crest  except  for  caudal  half  free  of  spots. 
Eggs  unknown.  Transformation  size  unknown.  Nevada. 

Rana  fishcri 
Plate  XIII-i 


GENERAL  ACCOUNT  67 

11.  Teeth  1/3  or  1/3 ;  four  to  six  rows  of  inner  papillae 
from  end  of  first  upper  row  to  end  of  lower  labial 
row;  a  row  of  heavy  inner  papillae  below  the 
third  lower  row  of  teeth;  spiracle  nearer  eye  than 
vent  i. 41-1,82.,  av.  1.62.;  nostril  nearer  eye  than 
snout,  1.0-1.42.,  av.  1.2.;  width  of  body  in  body 
length  1.45-1.86,  av.  1.6,  tadpole  large  (92.  mm.); 
second  upper  row  2.715  or  1/15  of  the  first  upper 
row  or  second  upper  row  absent,  belly  pale  chal- 
cedony yellow,  sulphur  yellow,  vinaceous,  pale 
grayish  vinaceous  or  vinaceous-buft";  upper  fail 
crest  with  a  black  line  or  row  of  large  black  spots,  more 
prominent  than  in  R.  grylio,  middle  of  musculature 
ivtth  another  black  line,  tail  dark  ivitb  pale  chalcedony 
yellow  spots.  Transformation  size  2.5-35  mm-  Eggs 
a  submerged  mass.  New  Jersey  to  Okefinokcc 
Swamp,  Ga.  Rana  vtrgattpes 

Plates  XI -13,  XIII  i 

dd    Papillary  border  on  side  of  labium  without  an  emargination,  tadpoles 
13-50  mm.  length;  labial  teeth  1/3  or  z/2..  HYLIDAE 

e.  Labial  teeth  z/i,  tip  of  tail  normally  black  (sometimes  lost),  median 
space  of  second  upper  tooth  row  wide  (like  Rana)  and  1.33  greater 
than  cither  lateral  portion,  horny  beak  1.1-1  35  m  first  upper  row, 
eye  dorsal  just  inside  the  lateral  outline  in  dorsal  aspect  (more  like 
Ranids).  Eye  i.o  1.66  (av.  i.zz)  nearer  tip  of  snout  than  spiracle; 
depth  of  tail  in  length  of  tail  3.2.5-5  o,  av.  40,  suborbital  region 
oblique,  not  vertical;  spiracle  to  eye  usually  equals  distance  from 
spiracle  to  vent;  spiracle  plainly  showing  from  dorsum,  sptracular  tube 
tn  Itfe  stands  out  at  an  angle  from  the  body  and  opening  tf  apart  from  the  body 
proper;  papillary  border  does  not  end  above  the  end  of  the  upper  row 
(like  Rana),  eggs  single,  occasionally  a  mass.  Acrts 

Plates  X-n;  XII  6,  XLV  4,  XLVI-9 

ee.  Labial  teeth  usually  1/3,  rarely  1/1,  tip  of  tail  not  normally  black, 
median  space  (sometimes  absent)  of  second  upper  tooth  row  narrow, 
1.6-4.0,  rarely  4  5-7.0  in  either  lateral  portion,  j  e.,  smaller  than 
cither  lateral  part,  horny  beak  or  mandible  1.3*  2..3  in  first  upper  row, 
eye  lateral  visible  from  ventral  as  well  as  the  dorsal  aspect;  eye  i.o- 
i  75  nearer  spiracle  than  tip  of  snout;  depth  of  tail  in  length  of  tail 
1.6-4  4>  av-  2-I~"3 '55  i  suborbital  region  vertical;  spiracular  tube  in  life 
parallel  with  opening  at  inner  edge  closely  connected  with  or  near  to 
body  proper.  HYLIDAE  exclusive  of  Acris 


68  HANDBOOK  OF  FROGS  AND  TOADS 

f.  Labial  teeth  occasionally,  or  frequently  1/1;  tadpoles  13-33  mm- 

g.  Tail  musculature  with  black-brown  lateral  band  with  light  area  below;  tail 
crests  clear  with  fine  elongate  fleckings,  single  row  of  papillae  on  lower 
labial  border  below  second  lower  row  of  teeth;  upper  fringe  somewhat 
angulate  in  middle  (like  H.  femoralis);  spiracle  1.4-1.9  nearer  base  of 
hind  legs  or  vent  than  tip  of  snout,  av.  1.41,  nostril  to  eye  1.37-1.0  in 
nostril  to  snout,  av.  1.6;  median  space  between  the  second  upper  row 
of  teeth  1^-5  in  either  lateral  portion;  ends  of  second  lateral  row  ex- 
tending not  at  all  or  slightly  beyond  the  end  of  the  upper  fringe;  horny 
beak  about  1.5-1.6  in  upper  fringe.  Eggs  an  irregular  mass. 

Pseudacns  nigrita  triseriata 

gg.  No  black-brown  dorsal  and  white  or  light  lower  band  on  tail  musculature;  tail 
fins  clear  except  where  heavily  pigmented  with  purplish  black  blotches 
on  the  outer  edge  of  the  tail  fin;  two  rows  of  papillae  on  lower  labial 
border  below  second  lower  row  of  teeth,  upper  fringe  not  perceptibly 
angulate  in  the  middle;  spiracle  1.0-1.6  nearer  base  of  hind  legs  or  vent 
than  tip  of  snout,  av  2..i6,  nostril  to  eye  i.o  in  nostril  to  snout; 
median  space  between  second  upper  row  of  teeth  1-3  in  either  lateral 
row;  ends  of  second  lateral  row  extending  beyond  upper  fringe  for  1/4 
to  1/6  of  either  lateral  portion,  horny  beak  1.75-1.15  in  length  from 
one  end  of  second  upper  lateral  row  to  end  of  other  lateral  row.  Eggs 
single.  Hyla  cructfer 

Plates  X-3,XII-i 

ff .  Labial  teeth  1/3 . 

g.  Third  row  of  labial  teeth  short,  shorter  than  horny  beak  or  0,10-0  40 
of  the  first  lower  row  in  length,  first  upper  row  very  angulate  (i  form), 
somewhat  angulate  (i  forms),  not  angulate  (3  forms),  no  flagellum 
ordinarily  present  yet  some  have  pointed  tails,  tadpoles  13-50  mm., 
light  papillary  development,  lower  labial  corner  not  with  three  or 
four  strong  rows  of  papillae,  one  or  two  rows  of  papillae  below  third 
lower  row  of  teeth  or  none  (Ji .  crucifer)',  the  papillae  extend  above  and 
beyond  the  end  of  the  upper  fringe  for  about  0.14-0.185  of  the  length 
of  the  upper  fringe. 

h.  Tadpoles  13-33  mm.,  eye  equidistant  between  spiracle  and  tip  of 
snout  (o.8--i.3  times);  spiracle  1.1-1.6  nearer  vent  or  base  of  hind 
legs  than  tip  of  snout,  spiracle  i  .0-3 .1  nearer  eye  than  vent,  papillae 
extend  above  the  fringe  for  0.16-0.15  of  the  length  of  the  first 
upper  row. 

i.  Musculature  with  no  distinct  brown  lateral  band  with  light  area  below; 
crests  usually  heavily  pigmented  with  purplish  black  blotches 
on  outer  rim,  nostril  to  eye  in  nostril  to  snout  2.  o,  spiracle  to 
vent  1.0-1.6  in  spiracle  to  tip  of  snout;  no  papillae  below  third 


GENERAL  ACCOUNT  69 

lower  labial  row  of  teeth  thus  appearing  as  a  goatee;  median  space  between 
second  upper  labial  row  1-3  in  either  lateral  portion;  horny  beak  in  first 
upper  row  1.75-1.15.  Eggs  single,  submerged.  End  of  second  upper  row 
extends  beyond  end  upper  fringe  1/4  to  1/6  width  of  second  lateral  row; 
upper  fringe  not  angulate  in  middle;  median  space  in  second  upper  row 
1.4-1.5  in  horny  beak;  third  lower  row  in  first  or  second  lower  row  3-5 
times;  second  lower  row  distinctly  longer.  Hyla  cructfer 

Plates  X-3,  XII -i 

ii.  Musculature  with  a  distinct  brown  lateral  band  above  with  light  area  below  >  i.e., 
bicolored;  crests  usually  clear  with  fine  scattered  fleckings,  sometimes  with 
fleckings  gathered  nearer  outer  rim,  one  or  two  rows  of  papillae  below 
third  lower  labial  row  of  teeth  (rarely  absent),  spiracle  to  vent  in  spiracle 
to  tip  of  snout  once  or  i.i  -i.o  or  1.1  times,  one  or  two  rows  of  papillae 
below  the  third  lower  labial  row  of  teeth;  median  space  between  second 
upper  row  1.6-5.0  times  in  either  lateral  portion,  horny  beak  in  upper 
fringe  1.3-1  3,  end  of  second  upper  lateral  row  to  end  of  first,  slightly  or 
distinctly  shorter,  or  rarely  slightly  beyond,  first  upper  row  very  angulate 
(i  form)  somewhat  angulate  (3  forms),  not  angulate  (2.  forms)  in  middle; 
median  space  (absent  in  i  form)  of  second  upper  row  1.5-3.5  *n  horny 
beak,  third  lower  row  of  teeth  from  i  1-4  times  in  first  or  second  lower 
row;  second  and  first  equal  (except  one  with  first  considerably  shorter 
and  one  equal  or  first  1.1-1.1  shorter).  Eggs  in  a  mass  except  P.  oculans 
where  single.  Pseudacris 

j.  Tadpoles  15^-13  mm.,  adults  11.5-17.5  mm  ,  eggs  single;  dorsal  crest 
to  vertical  midway  between  spiracle  and  eye,  spiracle  equidistant  from 
eye  and  vent;  median  space  3.5  in  horny  beak,  horny  beak  i.o  m  upper 
fringe,  horny  beak  i.o  in  first  or  second  lower  rows  of  teeth,  third 
lower  row  of  teeth  3-4  in  first  or  second  lower  rows;  third  lower  row 
4.0  in  first  upper  row,  first  upper  row  somewhat  angulate  in  middle; 
mouth  and  internasal  space  equal,  spiracle  1.7-1  i  nearer  vent  than 
snout,  one  row  of  papillae  below  the  third  lower  row  of  teeth;  dorsum 
of  body  tn  Itfe  with  definite  scattered  black  spots;  musculature  with  three 
bands,  apricot  buff  (light),  chestnut  brown  (dark),  martius  yellow 
(light).  Pseudacns  oculans 

Plates  X-8;  XII-8 

jj.  Tadpoles  2.3-36  mm.;  adults  19^-48  mm.,  eggs  in  a  mass,  dorsal  crest  to 
vertical  of  spiracle  or  within  i  or  z  mm.  of  such  vertical;  spiracle  to 
eye  1.0-3.0  or  3.1  nearer  eye  than  spiracle  to  vent  (one  form  sometimes 
equidistant);  median  space  usually  present,  1.5-3.0  in  horny  beak; 
horny  beak  1.3-1.3  in  first  upper  row;  horny  beak  in  first  or  second 


7o  HANDBOOK  OF  FROGS  AND  TOADS 

row  1111,  rarely  over  i.o  times,  third  lower  row  of  teeth  in  first  or  second 
lower  row  i  i- 1.8,  rarely  3.0  3.1  times,  third  lower  row  in  first  upper  row 
1.6  -3.8  occasionally  4  o.  Pstudacns  (exclusive  of  P.  oculans) 

(k.  Adults  15  -48  mm  ,  first  and  second  lower  tooth  rows  equal  and  united 
at  their  outer  ends,  median  space  of  first  upper  tooth  row  very  angulate; 
musculature  depth  3.0  in  tail  depth,  median  space  of  second  upper  row 
i. 6-i  7  in  either  lateral  part,  median  space  i  5  in  horny  beak,  horny 
beak  1.1  2.  3  in  first  upper  row,  third  row  1.3  in  horny  beak,  mtcrnasal 
space  i. 6 -i. 7  in  mterorbital  space,  tail  depth  greater,  brown  gray 
dorsally,  with  median  mterorbital  rectangular  darker  area  and  a  crescent 
at  rear  of  each  nostril,  belly  white.  (After  Bragg  ))  Pscudacns  stricken 

kk.  Adults  to  19  36  mm  ,  musculature  depth  in  tail  depth  i  1-1  9,  median 

space  of  upper  labial  tooth  row  somewhat  angulate  or  not  angulate, 

median  space  of  second  upper  row  1-5  in  cither  lateral  portion  (absent 

in  i  form,  sometimes  absent  in  another),  median  space  of  second  upper 

row  in  horny  beak  3  o  or  more,  horny  beak  i  3-2.  o  in  first  upper  row; 

third  row  13  3  o  in  horny  beak,  first  and  second  lower  rows  equal,  or 

first,  ri  i  i,  or  considerably  shorter,  free  at  ends  (in  one  form  almost 

united),  internasal  space  in  interorbital  1.33  i  4,  usually  i.o  or  more, 

rail  depth  in  tail  length  usually  11-40,  rarely  4  4. 

(1    Tail  depth  in  tail  length  1115,  musculature  of  tail  17  3.0  in  tail 

depth,  eye  to  spiracle  1.5  in  eye  to  tip  of  snout,  internasal  space  i  8 

in  mterorbital  space,  internasal  space  i.o  in  mouth,  nostril  equidistant 

from  eye  and  snout  or  i  15  nearer  eye  than  tip  of  snout,  iirst  upper 

row  not  angulate  in  middle,  median  space  of  second  upper  row  i  o- 

i  3  in  cither  lateral  part  or  3.0  in  horny  beak,  horny  beak  i  6-1  8  in 

first  upper  row,  or  i  6  in  first  or  second  lower  row,  third  lower  row 

1.5   i  6  in  horny  beak  or  16  1.8  in  first  or  second  row,  first  and  second 

lower  rows  equal  and  almost  united  on  their  ends,  black-brown  above, 

under  parts  bronzy  black,  black  along  side,  body  transparent.  (After 

Green  ))  Pseudacns  bracbypbona 

11.  Tail  depth  1.4  5.0,  seldom  under  i  5  in  tail  length,  musculature  1.1- 

i  6  in  tail  depth,  eye  to  spiracle  0.8  1.36  in  eye  to  snout,  internasal 

space  1.3  1.4  in  mterorbital  space,  internasal  space  about  equal  or  i.i 

or  i. i~i  7  in  mouth,  nostril  to  eye  1.1-1  o  in  nostril  to  snout. 

m.  Upper  fringe  of  teeth  not  angulate,  no  median  space  in  second 

upper  row  or  median  space  1.8-1.0  in  either  lateral  part  or  i.o  in 

horny  beak,  horny  beak  in  upper  first  row  of  teeth  or  fringe  1.3- 

1.7,  first  and  second  rows  unequal,  first  considerably  (1.1-1.5) 

shorter  than  second  row;  horny  beak  1.1-1.5  in  second  row  and 


GENERAL  ACCOUNT  71 

1.1-1.4  greater  than  first  row,  horny  beak  1.3-1  o  in  upper  fringe; 
third  lower  row  1.5-3.0  in  horny  beak;  one  row  of  papillae  under 
third  lower  row;  spiracle  nearer  eye  than  vent,  tail  depth  in  tail 
length  1.4,  musculature  of  tail  1.6  in  tail  depth,  third  lower  row 
1.0-1.3  in  first  lower  row,  1.3-3  1  in  second  lower  row,  gray  olive 
or  grayish  brown  tadpoles.  Pswdacrts  n.  clarkti 

Plate  XIII  4,  L-5 

mm.  Upper  fringe  of  teeth  somewhat  angulate,  median  space  of  second 

upper  row  1-7  in  cither  lateral  portion  of  this  row  (occasionally 

absent  in  P.  //.  ftnawtri),  or  3  or  more  in  horny  beak,  first  and 

second  lower  tooth  rows  equal  or  first  1.1-1.1  shorter  than  second 

row,  third  lower  row  1.3  1.8  m  horny  beak,  horny  beak  1.5  i.o 

in  upper  fringe  or  1.3-1.1  in  first  or  second  lower  rows;  spiracle 

to  eye  1.0-1.7  nearer  than  spiracle  to  vent;  tail  depth  15  3.15  in 

tail  length,  musculature  of  tail  1.1  1.5  in  depth  of  tail,  third 

lower  row  i  0-3.0  m  first  and  second  lower  row. 

n.  Two  rows  of  papillae  under  third  row  of  teeth,  median  spate 

of  second  upper  row  sometimes  absent,  if  present  ij  *j  or  3-7 

m  lateral  part,  horny  beak  i  o-i  i  in  first  or  second  lower 

rows  of  teeth,  third  lower  row  i  3-1.4  in  horny  beak  or  3  4 

m  upper  fringe.  Puudacns  n  fenarum 

Plate  XII  5 

nn.  One  row  of  papillae  under  third  lower  row  of  teeth;  median 
space  of  second  upper  row  i  5  in  either  lateral  row,  horny  beak 
i  3-  j  5  in  first  or  second  lower  rows  of  teeth,  third  lower  row 
i  8  in  horny  beak  or  3  o  in  upper  fringe. 

Ptcudacris  n  trtscnata 

[The  key  to  PtcuJacns  may  be  worthless,  and  the  characters 
given  here  may  be  extremely  variant.] 

hh.  Tadpoles  35-50  mm.,  tadpoles  not  bicolorcd  (brown  above  and  white 
below)  but  green,  citrine  tadpoles  or  scarlet-  or  orangc-uilcd  tadpoles; 
high-crested  tadpoles,  eye  1.0-1.75  nearer  spiracle  than  snout,  spiracle 
i  o-i  6  nearer  vent  than  tip  of  the  snout,  spiracle  i  15-1  5  nearer  eye 
than  vent  or  base  of  hind  legs,  papillae  extend  above  the  upper  fringe 
for  o  14-0  185  of  the  length  of  the  upper  fringe. 

i.  Tadpoles  50  mm.  in  length,  body  in  tail  1.3-3  15»  av-  M»  depth  of 
body  in  width  of  body  o  83-1.0,  av.  o  9,  depth  of  tail  10  14  mm  , 
beautiful  green  tadpoles,  young  tadpoles  with  a  black  saddle  spot  on 
the  back  of  the  musculature  near  its  base  and  with  a  light  line  from 
eye  to  tail,  one  row  of  papillae  below  lower  third  labial  row,  papillae 
extending  above  first  upper  row  for  0.15-0.185  of  the  fringe's  length, 


72  HANDBOOK  OF  FROGS  AND  TOADS 

dorsal  crest  extending  to  a  vertical  halfway  between  eye  and  spiracle. 
Eggs  single,  submerged.  North  Carolina  to  Louisiana.          Hyla  gratiosa 

Plates  X-5,6,  XII-i,  LXIX-5 


ii.  Tadpoles  35-45  mm.,  body  in  tail  1.1-1.0,  av.  1.6,  depth  of  body  in 
width  of  body  1.0-1.8;  depth  of  tail  5-9  mm.,  no  black  saddle  spot  in 
young  tadpoles. 

j.  Tadpole  small  (35  mm.),  dorsal  crest  extending  to  vertical  halfway 
between  spiracle  and  the  base  of  the  hind  legs;  depth  of  tail  in  tail 
length  2.. 5-3. 5,  av.  30,  nostril  to  eye  1.1-1 1  in  nostril  to  snout; 
mouth  in  interorbital  space  1.33-1.6,  mternasal  space  in  interorbital 
distance  i  33-1.1,  eye  just  touches  lateral  axis  or  is  below  it,  beak  in 
first  upper  row  1.5-1.7,  papillae  extending  beyond  the  end  of  the  upper 
row  0.15-0.185  of  the  length  of  the  upper  row,  two  rows  of  papillae 
below  third  lower  labial  row,  median  space  between  second  upper 
labial  row  1.15-10  in  either  lateral  portion;  third  lower  labial  row 
o.io  0.11  of  the  first  lower  row,  first  row  of  lower  labial  teeth  1.0-1.5 
times  the  horny  beak.  Eggs  strewn  in  water  among  sphagnum  (Noble 
and  Noble).  New  Jersey  to  South  Carolina.  Hyla  andtrsomt 

Plate  X-i,XII-3 

jj    Tadpole  medium  (40  and  45  mm  ),  dorsal  crest  extends  ahead  of 
spiracle  or  to  eye,  depth  of  tail  in  tail  length  i  5-3.1,  av.  1.75,  nostril 
to  eye  1.0-1.7  in  nostril  to  snout,  mouth  in  interorbital  space  i  4-1.0, 
internasal  in  interorbital  space  1.15-1.0,  eye  on  lateral  axis,  papillae 
extending  beyond  end  of  first  upper  row  1.4-1.5  of  the  length  of  the 
first  upper  row,  median  space  in  second  upper  labial  row  3-5  in  either 
lateral  portion,  third  labial  row  0.15-0.40  of  the  first  labial  lower  row; 
first  row  of  lower  labial  teeth  i  0-1.3  greater  than  the  horny  beak, 
k.  Dorsal  crest  to  the  vertical  halfway  between  spiracle  and  the  eye, 
depth  of  body  in  body  length  i  7-1.5,  musculature  of  tail  in  depth 
of  tail  1.75-1.4,  av.  1.9,  spiracle  1.4-1.3  nearer  eye  than  vent; 
mouth  1.0-1.4  larger  than  internasal  space,  av.  1.15,  two  rows  of 
papillae  below  the  third  lower  row  of  labial  teeth,  papillae  extend 
beyond  the  end  of  the  first  upper  row  0.11-0.15  of  the  length  of  this 
fringe,  horny  beak  in  upper  row  1.0^-1.3;  third  labial  lower  row 
0.15-0.40  the  length  of  the  first  lower  row.  Eggs  surface  or  sub- 
merged irregular  mass.  Eastern  Maryland  to  Florida  to  Texas  to 
Illinois.  Hyla  cmcrca 

Plates  X-i;  XII~4 


GENERAL  ACCOUNT 


73 


kk.  Dorsal  crest  extending  to  the  vertical  of  the  posterior  edge 
of  the  eye;  depth  of  body  in  body  width  1.5-1.0;  muscula- 
ture in  depth  of  tail  1.3-1.8,  av.  1.5,  spiracle  1.6-1.75  nearer 
eye  than  snout;  mouth  1.0-1.1  larger  than  internasal  space; 
one  row  of  papillae  below  the  third  lower  labial  row  of 
teeth,  papillae  extends  beyond  the  end  of  the  first  upper  row 
0.14-0.10  of  the  length  of  this  upper  fringe;  horny  beak  in 
upper  fringe  1.4-1.8;  third  labial  lower  row  0.15-0.33  the 
length  of  the  first  lower  row.  Eggs  loose  irregular  mass. 
Vancouver  to  Lower  California  and  east  to  Utah  and  Arizona. 

Hyla  regtlla 

gg.  Third  row  of  labial  teeth  long,  longer  than  horny  beak,  or  0.75-1.00  of 
the  first  lower  row  in  length;  first  upper  row  very  angulate  in  middle; 
flagellum  on  tail;  tadpoles  31-50  mm,,  heavy  papillary  development, 
lower  labial  corner  with  three  or  four  rows  of  papillae;  two  more  or  less 
complete  rows  of  papillae  below  third  row  of  teeth  (except  in  Hyla 
aremcolor),  papillae  extend  above  and  beyond  the  end  of  the  upper  fringe 
for  about  0.3-0.4  of  its  length. 

h.  Third  lower  labial  row  0.8-1.0  of  the  length  of  the  first  lower  row; 
dorsal  crest  extends  to  the  vertical  halfway  from  hind  legs  to  spiracle, 
to  spiracle,  or  halfway  from  spiracle  to  eye,  dorsal  crest  equal  to, 
greater,  or  less  than  depth  of  tail  musculature;  tadpoles  36-50  mm.; 
red  may  be  present  in  the  tail,  tail  crest  distinctly  or  more  or  less 
clear  of  spots  next  to  the  musculature,  tail  heavily  blotched  with 
dark  blotches  or  spots. 

i.  Median  space  between  lateral  upper  rows  5.0-10.0  times  in  either 
lateral  row,  spiracle  1.44-1.5  nearer  eye  than  vent,  width  of  body 
in  its  own  length  1.6-1.1,  eye  1.0-1.7  nearer  spiracle  than  tip  of 
snout;  tail  sometimes  suffused  with  coral  red  or  coral  pink, 
j.  Median  space  between  lateral  upper  rows  of  teeth  contained 
6.a-io.o  in  either  lateral  row,  first  and  second  lower  rows  of 
teeth  1.4-1.6  greater  than  horny  beak;  mouth  equal  to  inter- 
nasal  space;  depth  of  tail  in  length  of  tail  1.6-1.75,  av.  1.15; 
muscular  part  of  tail  in  depth  of  tail  1.8-1.3,  av.  1.1;  depth 
of  body  1.33-1.1  in  body  length,  av.  1.68;  dorsal  crest  usually 
equal  to  or  greater  than  musculature  depth;  center  of  belly 
solid  sulphur  yellow;  tail  )-j-banded;  light  lateral  band  bounded 
above  and  below  by  a  brown  band,  flagellum  clear  of  pigment; 
body  olivaceous  black.  Eggs  a  surface  film.  Virginia  to  Florida 
to  Texas .  Hyla  fcmorahs 

Plates  X-4;  XII-i3;  LXVIII-3 


74  HANDBOOK  OF  FROGS  AND  TOADS 

jj.  Median  space  between  lateral  upper  rows  contained  5.0-10.0  times 
in  either  lateral  row,  first  and  second  lower  rows  of  teeth  1.5-1.0 
greater  than  horny  beak;  third  lower  row  of  teeth  may  be  equal  to 
or  slightly  shorter  than  the  first  lower  row;  no  or  few  papillae  be- 
neath the  third  lower  row  of  teeth,  surely  not  a  complete  row; 
mouth  in  internasal  space  0.83-1.7,  av.  1.3;  depth  of  tail  in  length 
of  tail,  x. 85-5. 15,  av.  3 .75 ;  musculature  of  tail  in  depth  of  tail  1.18- 
1.76,  av.  1.6;  depth  of  body  in  body  length  1.75-1.3,  av.  i.o;  dorsal 
crest  halfway  to,  or  to,  vertical  of  the  spiracle;  dorsal  crest  less  than 
the  musculature;  flagellum  or  tail  tip  spotted;  body  greenish  olive 
or  deep  olive;  center  of  belly  solid  pale  cinnamon  pink.  Eggs  single, 
submerged  (Atsatt  and  Storer).  Western  Texas  (Devil's  River,  Fort 
Davis  Mountains,  etc.),  Utah,  California,  Mexico.  Hyla  arenicolor 

Plate  LX-6,7 

ii.  Median  space  between  lateral  upper  rows  contained  3.15-5.0  times  in 
either  lateral  row;  spiracle  1.11-1.5  nearer  eye  than  vent;  eye  about 
equidistant  between  spiracle  and  tip  of  snout;  internasal  space  in 
mouth  0.7-1.0,  dorsal  crest  extends  to  vertical  of  spiracle  or  halfway 
between  eye  and  spiracle;  dorsal  crest  equal  to  or  greater  than  muscu- 
lature depth;  muscular  part  of  tail  in  depth  of  tail  1.71-1  9,  av.  1.8; 
depth  of  tail  in  length  of  tail  3.1-4.0;  width  of  body  in  its  own  length 
i  .3-1 .7;  no  lateral  bands  in  tail;  tail  more  or  less  scarlet  or  orange  vermilion 
with  black  blotches  more  prominent  near  the  margins  of  the  crests;  bodies 
olive-green;  belly  conspicuously  white  or  very  light  cream.  Eggs  a 
surface  film.  Minnesota  to  Texas  to  Maine  to  Florida. 

Hyla  v.  versicolor 
Plates  X~9,io;  XII-n 

hh.  Third  lower  labial  row  0.75  of  the  length  of  the  first  row,  dorsal  crest 
extends  to  the  vertical  of  the  posterior  edge  of  the  eye;  dorsal  crest  usu- 
ally less  than  depth  of  the  musculature;  tadpoles  to  31  mm.;  width  of 
body  in  its  own  length  1.7-1.1,  av.  1.875;  depth  of  tail  in  length  of  tail 
1.1-3.3,  av.  1.8,  third  lower  row  of  teeth  not  equal  to  first  lower  row; 
median  space  between  lateral  upper  rows  contained  3.15-5.0  times  in 
length  of  either  lateral  row;  papillae  extend  above  and  beyond  the  ends 
of  the  upper  fringe  for  0.3-0.33  of  the  upper  fringe,  horny  beak  in  upper 
fringe  1.8-1.0;  no  bands  or  red  in  tail;  tail  crest  clear,  uniformly  sprinkled 
with  distinct  black  dots;  body  greenish  (like  H.  cinerea  or  H.  gratiosa); 
belly  testaceous  or  chalcedony  yellow.  Eggs  single,  submerged.  Texas 
to  Indiana  to  Florida  to  Virginia.  Hyla  squirella 

Plates  X-7;  XII-9 


GENERAL  ACCOUNT 


75 


Development  and  transformation: 

Some  frogs  have  limited  breeding  pc 
riods  and  other  species  may  breed  al- 
most any  month  in  the  year.  The 
males  usually  precede  the  females  to 
the  water  and  croak  vigorously  dur- 
ing breeding  time.  The  male  with  ils 
forearms  seizes  the  female.  In  almost 
all  frogs  the  eggs  are  fertilized  just  at 
or  slightly  after  the  extrusion  of  the 
eggs.  At  first  no  envelopes  about  the 
eggs  are  apparent,  and  the  egg  mass 
may  feel  soft  and  sticky.  After  a  few 
minutes  this  substance  absorbs  water, 
and  each  egg  is  then  revealed  with  its 
vitelline  membrane  and  one  or  more 
jelly  envelopes. 

The  eggs  hatch  in  3  to  25  days, 
depending  on  temperature  and  other 
conditions.  At  hatching  the  larva  has 
a  distinct  neck,  with  a  prominent 
head  and  body.  The  tail  is  very  small 
or  absent.  On  the  ventral  side  of  the 
head  is  an  invagination  or  depression 
which  is  to  be  the  mouth.  Behind  this 
comes  the  ventral  adhesive  disk  or 
disks,  which  help  the  little  creature 
to  attach  itself  to  the  egg  mass  or  to 
hang  itself  upon  some  plant.  In  front 
of  the  mouth  are  two  deep,  dark  pits 
which  later  become  the  nostrils.  On 
either  side  of  the  head  appear  swell- 
ings which  become  the  external  gills, 
The  eyes  do  not  yet  appear. 

As  development  goes  on,  the  exter- 
nal gills  appear  as  branched  organs: 
two  or  three  on  a  side;  the  eye  shows 
as  a  ring  beneath  the  skin;  and  the 
tail  grows  and  presents  a  middle  mus- 
cular portion  where  the  muscle  seg- 
ments clearly  show.  This  middle  part 
supports  a  thin,  waferlike  tail  fin  the 
parts  of  which  are  called  the  lower 


?    I 


Plate  XV.  Development  of  the  gopher 
frog,  Rana  capita,  i.  Egg  mass  (X%)-  2. 
Eggs  (X^)-  3-7-  Tadpoles  (X%)*  U- 
Lateral  lines  and  spiracle;  5,  with  two  legs; 
6,  with  three  legs;  7,  with  four  legs.]  8. 
Transformed  frog  (X%)«  9-  Adult 


76  HANDBOOK  OF  FROGS  AND  TOADS 

and  upper  crests.  The  nasal  pit  shifts  in  position  and  becomes  the  nostril,  and 
the  vent  opens.  The  mouth  appears,  and  dependence  on  the  yolk  of  the  belly 
ceases.  Soon  the  external  gills  begin  to  disappear,  a  lateral  dap  or  fold  of  skin 
connects  the  head  with  the  body,  and  the  neck  region  disappears.  Beneath 
this  fold  internal  gills  develop.  Usually  on  the  left  side  but  on  the  middle  line 
in  the  belly  in  ribbed  frogs  and  narrow-mouthed  toads,  the  flap  does  not  close 
completely,  but  leaves  an  opening,  the  spiracle.  The  water  passes  into  the 
mouth  over  the  internal  gills  and  out  of  this  hole.  On  the  mouth  a  membra- 
nous, fringed  lip,  with  upper  and  lower  portions  (labia)  comes  into  being. 
At  the  portal  are  horny  jaws  or  mandibles.  On  the  upper  and  lower  portions 
are  ridges  of  horny  teeth.  The  eyes  are  no  longer  covered,  pigmented  rings  but 
are  now  at  the  surface.  The  intestine  has  become  much  elongated  and  coiled 
and  in  some  can  be  seen  through  the  skin.  The  skin  of  the  back  and  head 
comes  to  have  a  series  of  sense  organs  (or  lateral  line  dots). 

The  buds  of  the  hind  limbs  begin  to  appear.  The  fore  limbs  start  to  develop 
beneath  the  skin.  When  the  hind  limbs  have  reached  considerable  size,  the 
left  arm  comes  out  through  the  spiracle,  or  the  skin  breaks  down  and  the  right 
arm  appears  through  the  skin  or  the  skin  weakens  for  its  egress.  It  is  held  that 
the  kft  arm  normally  comes  out  first,  but  often  the  right  arm  appears  first. 

The  process  of  transformation  continues.  The  tail  crests  decrease  in  size  and 
the  creature  begins  to  live  on  its  tail — that  is,  to  absorb  it.  The  gills  vanish,  and 
the  lungs  begin  to  serve  as  the  sole  respiratory  organs,  if  the  skin  be  not  con- 
sidered. The  tadpole  appears  more  and  more  at  the  surface  or  near  the  shore. 
The  eye  assumes  eyelids.  The  tadpole  mouth  fringe,  with  its  horny  jaws  and 
horny  teeth,  is  discarded,  and  a  true  frog  mouth  begins  to  appear.  The  long 
intestine  becomes  wonderfully  shortened,  for  a  carnivorous  diet,  and  the  small 
frog,  with  a  vestige  of  a  tail,  is  ready  to  leave  the  water.  This  process  is  termed 
transformation  or  metamorphosis. 

In  the  accompanying  table  of  51  tadpoles,  wherein  total  length,  tail  length, 
body  length,  and  range  of  transformation  size  appear,  we  have  a  surprising 
correlation  in  an  ascending  scale  for  all  four  characters.  The  first  criterion  of 
arrangement  is  the  arbitrary  choice  of  total  length.  Strictly  speaking,  body 
length  and  transformation  sizes  should  have  governed  the  arrangement,  but 
all  seem  more  or  less  correlated.  The  most  striking  point  is  that  the  body 
length  of  the  largest  tadpoles  for  each  species  falls  within  the  range  of  trans- 
formation size  in  all  except  six  forms.  In  two  of  these  the  differences  are  only 
2  mm.  and  i  mm.  In  the  spadefoot,  Scaphiopus  couchii,  the  body  length  of  the 
tadpole  is  2  mm.  greater  than  the  known  size  at  transformation,  and  in  Ham- 
mond's spadefoot,  Scaphiopus  hammondii,  it  is  5  mm.  greater  than  the  known 
transformation  size.  In  Scaphiopus  holbroo^ii  no  such  condition  obtains.  Is 
it  because  these  two  species  are  desert  species  which  breed  in  the  most  transi- 
tory places,  and  is  there  consequently  a  decided  shrinking  in  body,  before 
transformation,  to  speed  development?  Many  have  thought  there  is  consid- 


GENERAL  ACCOUNT 


77 


erable  shrinkage  in  the  body  of  the  tadpole  before  transformation.  This  may 
be  true  of  some  species,  but  in  the  following  table  there  is  no  striking  decrease 
as  in  the  paradoxical  toad  (Pseudis  paradoxus).  Rather,  the  uniform  parallel 
in  body  length  of  tadpole  and  of  transformed  individual  is  surprising  even  to 
the  present  authors,  who  have  long  worked  with  them. 

At  least  35  of  the  51  tadpoles  arc  less  than  50  mm.  in  total  length,  and  of  the 
26  species  whose  tadpoles  are  unknown  or  not  minutely  described  at  least  12 
more  will  doubtless  fall  in  this  group  of  small  or  medium  tadpoles.  In  our 
descriptions  we  have  used  small,  medium,  and  large  for  tadpoles.  We  have 
called  small  25-35  mm«;  medium  40-50  or  55  mm.;  large  60-140  mm.  (large, 
60-86;  quite  large,  95-100;  very  large,  110-145). 

In  the  table  at  least  32  species  have  tadpoles  with  body  lengths  of  18  mm.  or 
less.  Probably  in  time  we  will  know  that  50  of  our  species  fall  in  this  class,  or, 
put  in  another  way,  at  least  two-thirds  of  our  frog  species  will  be  found  to 
transform  at  18  or  20  mm.  or  less.  Kxcept  for  the  true  frogs  (Ranidae),  Ham- 
mond's spadcfoot,  and  a  toad  or  two  (Bufo),  no  species  grows  tadpoles  over 
2  inches  (50  mm.)  in  length. 

We  cannot  say  that  the  order  of  maximum  length  of  tadpoles  corresponds 
with  the  order  of  maximum  length  of  adults.  As  would  be  expected,  the  small- 
est swamp  cricket  frogs  and  narrow-mouthed  toads  have  small  tadpoles,  but 
with  them  appear  most  of  the  toads  (Bufo)  and  spadefoots  (Scaphiopus  ex- 
cept S.  hammondti).  Toads  and  spadefoots  arc  much  larger  than  the  tree  toads 
(Hyla  and  Acns),  which  follow  with  larger  tadpoles.  Beginning  with  the  tree 
frogs  through  the  true  frogs  (Ramdac),  the  order  of  tadpole  length  more  or 
less  corresponds  to  that  of  adult  size. 

Greatest      Greatest        Trans- 
Greatest  tail  body          forma- 
Name                                     length          length         length         lion  size 


Bufo  qucrdcus 
Bufo  debilis 
Pseudaci  is  oc  ulat  is 
Pseudacris  mgnta  trisenata 
Pseudacris  n.  septenttionalts 
Bufo  valliceps 
Bufo  terrestris 
Scaphiopus  couchii 
Pseudacris  brachyphona 
Microhyla  olivacea 
Bufo  punctatus 
Microhyla  carolinensis 
Scaphiopus  h.  hurterii 
Bufo  canorus 
Bufo  w.  jowleri 


mm. 


22 


23 

24 
24.5 

25 

25 

26 

26.4 

27 
27 
27 


mm. 


14.2 
14.2 

13 

14.2 

ii 

15.4 
16 
16.4 
16 


mm. 


8.8 
8 

10 

98 


10 
IO 
10 
12 

10 


mm. 
7-8 
8-1 1 

7-9 
7.5-11 

7-5-13 

7.5-12 

6.5-10.5 

7.5-12.5 

8 

10-12 

9-1 1 
7-12 
8-10 
10.5 


78 


HANDBOOK  OF  FROGS  AND  TOADS 


Greatest 
Name                                    length 

Greatest 
tail 
length 

Greatest 
body 
length 

Trans- 
forma- 
tion size 

mm. 

mm. 

mm. 

mm. 

Bufo  a.  americanus 

27.8 

17.6 

IO.2 

7-12 

Bufo  w.  woodhousii 

27 

*7 

10 

10-17 

Scaphiopus  h.  holbroolyi 

28 

16.4 

12 

8.5-12 

Bufo  compactilis 

28 

16 

12 

12 

Bufo  cognatus 

29 

i7 

12 

Pseudacrls  n.  clartyi 

30 

*9 

II 

8-I3 

Hypopachus  cuncus 

3° 

10-12 

Bufo  hemiophrys 

9-13-5 

Hyla  squlrella 

3* 

20 

12 

11-13 

Pseudacrls  ornata 

32 

18 

14 

I4-l6 

Leptodactylus  labialis 

32 

15 

17 

16 

Pseudacrls  n.  feriarum 

33 

22.4 

10.6 

8-12 

Hyla  crucifer 

33 

22 

II 

9-14 

Hyla  fern  or  alts 

33 

22.5 

IO 

10-15 

Pseudacris  strecl(erl 

33 

21 

12 

Hyla  andersonil 

35 

23 

12 

11-15 

Bufo  exsul 

35 

2O 

15 

Hyla  c.  cinerea 

40 

25 

15 

12-17 

Acrls  g.  gryttus 

42.2 

29.2 

J3 

9-15 

Hyla  wrlghtorum 

43 

26 

17 

11-17 

Bufo  b.  boreas 

44 

9.5-12 

Hyla  v.  verslcolor 

46.6 

30.8 

15.8 

13-20 

Hyla  regilla 

46.6 

3° 

16 

11-17 

Acris  g.  crepitans 

47 

32 

15 

11-15 

Rana  b.  boylii 

47 

29 

18 

23-30 

Hyla  gratiosa 

5o 

3i 

*9 

14-20 

Hyla  arenlcolor 

5° 

33 

15-4 

14-16 

Rana  a.  aurora 

5° 

3° 

20 

17-21 

Rana  s.  sylvatlca 

50 

33 

»7 

16-22 

Rana  s.  cantabrlgensls 

50 

3« 

20 

14-22 

Ascaphus  truei 

5* 

33 

18 

14-18 

Hyla  baudlnii 

21 

Scaphiopus  hammondii  Intermontanus 

52 

3« 

22 

20-24 

Bufo  b.  halophllus 

56 

33 

23 

12-15 

Bufo  b.  nelsoni 

I5-l8 

Rana  p.  sphenocephala 

62.5 

37-5 

25 

18-33 

Scaphiopus  h.  bombtfrons 

63-95 

Scaphiopus  h.  hammondii 

65 

37 

28 

13-23 

Rana  p.  pretiosa 

68.5 

43 

25.5 

16-23 

Rana  cascadae 

70 

22 

13-17 

Rana  b,  muscosa 

70 

20-24 

Cj 

Name 

ffcNfcKAL  ACCOl 

Greatest 
length 

JNT 

Greatest      Greatest 
tail             body 
length         length 

79 

Trans- 
forma- 
tion size 

mm. 

mm.           mm. 

mm. 

Rana  b.  sierrae 

72 

48              23.5 

21-27 

Rana  palustris 

75.8 

48.8            27 

19-27 

Rana  aurora  draytonii 

83 

55              ^8 

27 

Rana  fisheri 

83 

28-30 

Rana  areolata  circulosa 

30 

Rana  capita 

81 

53             29 

27-35 

Rana  pipiens 

84 

56             28 

18-31 

Rana  damitans 

84.8 

57             27.8 

28-38 

Rana  tarahumarae 

88 

56             32 

3°-35 

Rana  virgatipes 

92 

63             30 

23-31 

Rana  septentrionalis 

99 

67             32 

29-40 

Rana  hccfacheri 

95 

53-5           4i-5 

3i-49 

Rana  grylio 

no 

64.4          35.6 

32-49 

Rana  catesbeiana 

142 

97             45 

43-59 

Bufo  alvarius 

Not  described 

Bufo  californicus 

Not  described 

Bufo  insidior 

Not  described 

Hyla  avivoca 

Not  described 

Hyla  septentrionalis 

Not  described 

Pseudacris  brimlcyi 

Not  described 

Pseudacris  n.  nigrita 

Not  described 

Pseudacris  n.  vermtosa 

Not  described 

Rana  a.  aicolata 

Not  described 

Rana  scvosa 

Not  described 

Rana  onca 

Not  described 

Journal  notes:  These  arc  customarily  from  our  field  notes.  They  treat  of 
habitats,  general  habits,  or  breeding;  sometimes  of  experiences  ir  collecting  the 
frogs;  of  their  enemies;  or  of  their  usefulness  and  associated  ecological  fea- 
tures. Occasionally,  you  will  find  comparisons  with  closely  related  forms. 
Under  topics  such  as  "Voice,"  excerpts  from  our  field  notes  are  included.  In 
1925-1934  we  worked  on  Texan  forms,  and  in  the  first  edition  of  this  hand- 
book only  summaries  of  these  forms  appeared.  In  the  present  edition  more 
notes  are  added  on  the  southwestern  and  western  forms.  These  are  based 
mainly  on  a  recent  (1942)  25,ooo-mile  trip  in  that  region. 

Authorities'  corner:  Here  are  given  references  to,  or  quotations  from,  some 
of  the  more  important  works  on  the  various  frogs.  We  would  have  liked  to 
publish  excerpts  from  all  the  authors  cited,  but  space  limitations  forbade.  Pa- 
pers that  have  appeared  since  the  completion  of  the  manuscript  (1947-1948) 
have  also  been  listed.  These  references  are  in  no  sense  a  bibliography  of  the 
form. 


8o 


HANDBOOK  OF  FROGS  AND  TOADS 


Figure  i.  American  bell  toad,  Ascaphidae:  Ascaphus  truel.  i.  Short  anal  tube  of  female. 
2.  Light  band  across  head.  3.  Rear  of  femur.  4.  Tail  of  male,  extending  from  ventral  side 
of  body.  5.  Vent.  6.  Horny  excrescences  (secondary  breeding  characters)  of  the  male. 


Figure  2.  Spadefoots,  Scaphiopodidae:  Scaphiopus.  i.  Wide  interorbital  space.  2.  Upper 
eyelid.  3.  Small  round  parotoids.  4.  Fleshy  webs.  5.  Broad  waist.  6.  Pectoral  glands.  7. 
Vertical  pupil.  8.  Two  metatarsal  tubercles,  the  outer  large  and  with  a  cutting  edge. 


Keys 

FAMILIES 


A.  Male  with  taillikc  process,  female  with  short  anal  tube,  no  tympanum; 
pupil  elliptically  vertical;  upper  jaw  toothed;  short  ribs  present;  inter- 
nasal  space  4.66-8.0  in  L  ;  light  band  across  head;  size  small,  i  1/8-1 
in.  (2.8-51  mm.).  Ribbed  toads,  ASCAPHIDAE 

Plate  XVI 

AA.  Male  without  tail;  female  without  short  anal  tube;  ribs  absent;  inter- 
nasal  space  7.77-11.4  in  L. 
B.  Waist  wide,  body  broad  and  thick;  hind  limbs  short. 

C.  With  transverse  fold  of  skin  across  head  behind  eyes,  size  small, 
3/4-1  5/8  in.  (19-41  mm.);  no  tympanum;  no  parotoids;  snout 
pointed,  head  narrow,  fingers  and  toes  without  webs  except 
slight  in  Hypopachus;  eyes  small  and  depressed. 

Narrow-mouthed  toads,  BREVICIPITIDAE 
Plates  CXXIV--CXXVI 

CC.  Without  transverse  fold  of  skin  across  head,  behind  eyes;  size 
medium  to  large,  i  1/1-8  4/5  in.  (37-110  mm.),  except  B.  debtlis 
and  B.  quercicus  (1/3  in.),  tympanum  distinct  or  indistinct; 
parotoids  present  except  in  i  species  of  Scaphiopus;  feet  with 
extensive  fleshy  webs;  snout  blunt,  eyes  large;  head  broad. 
D.  Pupil  vertical  (by  day);  parotoids  absent  in  i  species,  present 
but  rounded  and  indistinct  in  i  species,  sole  without  sub- 
articular  tubercles,  skin  relatively  smooth;  venter  smooth; 
no  cranial  crests;  males  without  discolored  throats. 

Spadefoots,  SCAPHIOPODIDAE 
Plates  XVII-XXIII 

DD.  Pupil  not  vertical,  parotoids  present  and  elevated;  sole  with 
subarticular  tubercles;  skin  warty,  venter  usually  granulated; 
cranial  crests  present  in  most  species,  lacking  in  Bufo  boreas, 
B.  canorus,  B.  compacttlis,  B.  dcbtlis,  5.  cxsul,  B.  insidior,  B. 
punctatus;  males  usually  with  discolored  throats. 

Toads,  BUFONIDAE 
Plates  XXIV-XLIV 

BB.  Waist  narrow,  body  narrower  and  thinner;  hind  limbs  long;  no 
parotoid. 

C.  Disks  on  digits,  neither  thumb  nor  other  fingers  enlarged  in  male. 
D.  Disks  transverse;  venter  usually  smooth,  subarticular  tuber- 
cles saw-toothed;  eggs  large,  male  throat  not  discolored. 

Robber  frogs,  LEPTODACTYLIDAE 
(Syrrhophus,  Eleuthtrodactylus) 
Plates  LXXVII-LXXXIV 


82  HANDBOOK  OF  FROGS  AND  TOADS 

DD.  Disks  round,  large  or  small;  pupil  elliptically  horizontal;  sub- 
articular  tubercles  rounded,  venter  usually  granular  or  areolate; 
male  throat  discolored.  Tree  frogs,  HYLIDAE 

Plates  XLV-LXXVI 
CC.  No  disks  on  digits. 

D.  Extensive  webs  on  toes,  thumb  of  male  enlarged  at  base; 
venter  smooth.  Frogs,  RANIDAE 

Plates  LXXXV-CXXIII 

DD.  Toes  free;  males,  if  different,  with  strong  conical  spine  on  inner 
side  of  first  digit  and  another  on  each  side  of  breast,  and 
forehmb  much  enlarged. 

Robber  frogs,  LEPTODACTYLIDAE  (Lcptodactylus,  etc.) 
Plates  LXXXI,  LXXXII 

Spadefoots,  SCAPHIOPODIDAE:  Scaphiopus 

A.  Parotoid  absent  or  indistinct,  tympanum  indistinct,  no  pectoral  gland. 

B.  Hind  limb  longer  (0.78-0.96  in  L.);  fore  limb  longer  (1.51-1.19  in 

L.);  fourth  finger  longer  (6.18-8.0  in  L  ),  foot  with  tarsus  longer 

(1.57-1.76  in  L),  fourth  toe  longer  (1.66-3.2.9  in  L.);  size  small, 

i  1/1-1 1/5  in.  (37-61  mm.);  back  uniform  or  with  light  bands. 

Hammond's  spadefoot,  Scaphiopus  hammondii 
Plates  XVHI-XX 

Limits  of  range  and  gradations  need  to  be  worked 
out.  After  Tanner  we  present  this  synopsis: 

(C.  "Interorbital  boss  present,  rounded,  not  elongate  tubercle,  fronto- 
parietal  fontanelle  present,  body  smooth  or  less  rugose  dorsally 
than  CC;  head  width  narrower"  (width  of  head  in  L.  1.6-1.95, 
av.  1.74,  eye  smaller,  10.1-11.4  in  L.-- A.H.W.). 

Scaphtopus  h.  bombtfrons 
Plate  XIX 

CC.  "No  intcrorbital  boss  present.  In  some  specimens  of  intermontanus 
there  is  a  glandular  interorbital  elevation  which  resembles  the 
true  boss  in  bombtfrons"  Head  wider  (width  of  head  in  L.  1.11- 
1.58,  av.  i  35;  eye  larger,  7.3-8.6  in  L. — A.H.W.). 
D.  "Body  rugose  or  with  many  individual  prominences  or  warts." 
E.  "No  fronto-parietal  fontanelle;  interorbital  space  with 
prominent  fronto-parietal  bones  forming  ridges  ...  or  in 
some  specimens  the  interorbital  space  is  filled  with  a  glan- 
dular prominence  resembling  the  bombtfrons  species,  confined 
in  the  main  to  the  Great  Basin  area."  (Hind  toes  longer, 
second  toe  4.3-6.1  in  L.,  third  toe  3.1-4.0  in  L.;  fourth 
toe  1.1-1.66  in  L. — A.H.W.)       Scaphiopus  h.  tntcrmontanus 

Plate  XX 


KEYS  83 

EE.  "A  fronto-parietal  fontanellc  present;  interorbital  space 
smooth;  size  intermediate  between  bombtfrons  and  inttrmon- 
montanus;  .  .  .  found  on  Pacific  Coast  south  into  Arizona  and 
Texas."  (Hind  toes  shorter,  second  toe  6.1-7.3  in  L.,  third 
toe  4.0-4.1  in  L.,  fourth  toe  1.6-3.0  in  L. — A.H.W.) 

Scaphtopus  h.  hammondn 
Plate  XVIII) 

BB.  Hind  limb  shorter  (0.89-1.15  in  L  ),  fore  limb  shorter  (1.0-1.31  in 
L  ),  fourth  finger  shorter  (8  O-IT  i  in  L.),  foot  with  tarsus  shorter 
(1.7-135  in  L);  fourth  toe  shorter  (3.01-4.0  in  L.);  size  larger, 
i  7/8-3  1/5  m.  (48-80  mm.);  back  greenish,  more  or  less  marbled 
with  light.  Couch's  spadcfoot,  Scaphiopus  couchit 

Plate  XVII 

AA.  Tympanum  distinct,  parotoid  distinct,  pectoral  glands  present,  size 
1-17/8  in.  (50-71  mm.).  Scapbtopus  holbrookit 

Plates  XXI-XXIII 

B.  Head  to  angle  of  mouth  smaller  (3.14-3.81  in  L.),  width  of  head 
smaller  (1.58-1.66  in  L.),  snout  smaller  (5.4-6.3  in  L.),  tympanum 
smaller  (11.5-16.0  in  L  ). 

Hurter's  spadefoot,  Scapbiopus  holbrookit  hurtcnt 
Plate  XXIII 

BB.  Head  to  angle  of  mouth  greater  (1.93-  3.56  in  L.);  width  of  head 
greater  (i  14-1  75  in  L.),  snout  larger  (5.17  -6.6  in  L.);  tympanum 
greater  (10.1-11  o  in  L  );  skin  relatively  smooth  with  two  or  more 
evident  light  dorsal  stripes.  Spadefoot,  Scaphiopus  h.  holbrooktt 

Plate  XXI 

(C.  * 'Great  amount  of  white  on  back,  ilanks,  and  upper  surface  of 
limbs,  vermiculated  irregular  white  bands." 

Key  West  spadcfoot,  Scaphtopus  holbrookti  albtts 
Plate  XXII) 

Toads,  BUFONIDAE.  Bufo 

A.  Gland  on  leg,  fold  skin  on  tarsus,  warts  at  angle  of  mouth. 

B.  Crests  curved  around  rear  of  eye,  size  large,  3  1/5-6  3/5  in.  (8a- 
165  mm.),  skin  smooth;  head  broad  (1.1-1  8  in  L.),  color  uniform, 
glands  conspicuous  on  both  tibia  and  femur,  tympanum  medium 
(11-15  m  L.).  Colorado  River  toad,  Bufo  alvarius 

Plate  XXIV 

BB.  Crests  absent,  gland  on  tibia  only,  pitted  warts  on  back,  tympanum 
small  (14-16  in  L.). 


84  HANDBOOK  OF  FROGS  AND  TOADS 

C.  Dimorphic;  interparotoid  interval  less  than  width  of  gland;  parotoid 
and  leg  gland  obscured  by  pattern;  skin  smooth;  male  uniform  green; 
female  spotted;  size  small,  1-3  in.  (50-75  mm.)- 

Yosemite  toad,  Bufo  canorus 
Plate  XXXI 

CC.  Not  dimorphic;  parotoids  widely  separated,  interval  greater  than 
width  of  gland;  parotoid  and  leg  glands  evident,  size  small,  medium 
or  large  (41.5-115  mm.)  i  4/5-5  in.  Bufo  boreas  group 

D.  Eyelid  narrower  (11.8-14.6  in  L);  eye  smaller  (94-11.7  in  L.), 
head  narrower  (i  66-3.1  in  L.),  third  toe  longer  (1.8-3.5  m  L  )i 
fourth  toe  longer  (1.0-1.5  *n  L.)-  "Spread  of  hind  foot  from  end 
of  first  toe  to  the  fifth  toe  more  than  36%  of  length"  (Camp,  1917^. 

Northwestern  toad,  Bufo  boreas  boreas 

Plate  XXVII 

DD.  Eyelid  wider  (7.3-14  in  L.),  eye  larger  (7.0-10.5  in  L.);  head  wider 

(1.4-3.0  in  L.),  third  toe  shorter  (3.1-4  15  in  L.),  fourth  toe  shorter 

(i  53-3.8  in  L.),  spread  of  hind  foot  less  than  36%  of  the  length. 

E.  Dorsum  almost  entirely  black,  size  small,  i  4/5-1 1/1  in.  (44- 

61.5  mm  ),  eyelid  wider  (7  3-11  in  L  ),  width  of  head  1.77-3.0 

in  L.,  fourth  toe  longer  (1.53-1.66  in  L.),  venter  heavily  spotted 

with  black.  Black  toad,  Bufo  exsul 

Plate  XXXV 

EE.  Dorsum  gray,  green,  or  brown  spotted,  size  medium  to  large, 
i  4/5-5  in.  (41  5-115  mm.),  eyelid  narrower  (9.3-14  in  L.), 
fourth  toe  shorter  (1.57-3.8  in  L),  venter  uniform  white  or 
lightly  or  moderately  spotted. 

F.  Size  medium  to  large,  1 1/5-5  *n-  (60-115  mm.),  hind  limb 
longer  (0.8-1  o  in  L.);  head  wider  (1.4-1.85  in  L.),  fourth 
toe  shorter  (3.3-3  8  in  L.),  third  toe  longer  (3.1-3  8  in  L  ). 

California  toad,  Bufo  boreas  balophtlus 
Plate  XXVIII 

FF.  Size  small  to  medium,  i  4/5  to  1 1/1  in.  (41.5-61  mm.), 
hind  limb  shorter  (1.0-1.1  in  L.),  fore  limb  shorter  (1.86- 
1.15  in  L.),  head  narrower  (1.8-3  i  m  L  )»  fourth  toe  longer 
(1.57-1.94),  third  toe  shorter  (3.5-4.15  in  L.),  "elbows 
and  knees  not  meeting  when  appressed  to  the  sides  of  the 
body"  (Stejneger).  Amargosa  toad,  Bufo  b.  nelsoni 

Plate  XXIX 


KEYS  85 

AA.  No  gland  on  leg;  no  fold  of  skin  on  tarsus. 

B.  Femur  almost  entirely  enclosed  in  body  skin;  vocal  sac  elliptical 
(sausage),  i  sole  (metatarsal)  tubercles  with  cutting  edge. 
C  Crests  prominent;  boss  on  snout;  interorbital  narrow  (11-19  *n 
L.) — less  than  internasal,  middorsal  stripe  and  light-bordered 
large  dark  spots  (sometimes  small  spots). 

Great  Plains  toad,  Bufo  cognates 
Plate  XXXII 

CC.  Crests  absent,  interorbital  broad  (9.7-11.7  in  L.») — greater  than 
internasal;  drab  with  small,  dull  citrine  spots 

Spadefoot  toad,  Bufo  compactilis 
Plate  XXXIII 

BB.  Half  or  more  of  femur  free  from  body  skin,  outer  metatarsal  tubercle 
without  cutting  edge. 

C.  Parotoids  oval  to  elongate  (sometimes  triangular  in  Bufo  qucr- 
ctcus). 

D.  Crests  absent  or  obscure,  parotoids  broadly  oval,  divergent, 
interorbital  broad  (9.4-11  8  in  L.),  narrowing  forward; 
snout  7.0-8.15  in  L. 

Southern  California  toad,  Bufo  calif ornicus 
Plate  XXX 
DD    Crests  present. 

E.  Size  small,  3/4  i  1/4  m.  (19  31  mm.),  vocal  sac  a  sausage; 
mouth  small  (4.0  4  6  in  L  ),  tympanum  small  (14-  10  in 
L.),  snout  long  (5.7-7  o  in  L  ),  yellow  stripe  down  mid- 
back,  many  red  tubercles.  Oak  toad,  Bufo  qucmcus 

Plate  XL 

EE.  Size  greater,  i  5/8-4  3/4  in.  (40-118  mm  ),  snout  shorter 
(6.1-9.4  in  L  ). 

F.  Mouth  small  (4.1^4  7  in  L  ),  tympanum  small  (14-11 
in  L.),  head  narrower  (i  5-3.1  in  L  );  head  shorter 
(3.4-3.9  in  L.),  crests,  a  boss,  from  snout  to  rear  of 
eye,  with  sides  parallel,  snout  7.4-9  3  in  L. 

Canadian  toad,  Bufo  hcmiophrys 
Plate  XXXVI 

FF.  Mouth  large  (3.3-4.3  in  L.),  tympanum  large  (10- 
18.1  in  L  ),  head  wider  (1.3  in  L.). 
G.  Crests  prominent  with  knobs  in  rear,  skin  finely 
and   evenly   roughened   with   tubercles   between 
larger  warts,  red,  gray,  or  black;  snout  6  -8  in  L. 
Southern  toad,  Bufo  tcmstris 
Plate  XLI 


86  HANDBOOK  OF  FROGS  AND  TOADS 

GG.  Crests  low,  paired  spots  of  darker  color  down  back,  super- 
ciliary crests  meeting  postorbital  at  right  angles,  snout 
6.1-9.4  in  L. 

H.  Small  uniform  warts  on  back;  several  warts  in  each  dark 
dorsal  spot;  dark  pectoral  spot,  no  preparotoid  longitu- 
dinal crests,  under  parts  usually  unspotted;  back  greenish, 
size  smaller,  1-3 J^  in.  (51-  81  mm.);  snout  7.7-9.4  in  L. 
Fowler's  toad,  Bufo  woodhousn  fowleri 
Plate  XLIV 

HH.  Large  dorsal  warts;  one  or  two  warts  in  each  dark  dorsal 
spot,  under  parts  spotted  or  plain. 

I.  Many  warts  spiny,  particularly  on  hind  limbs,  paro- 
toids  parallel,  closest  together  at  midpoint;  parotoid 
on  dorsolateral  line  or  on  dorsum;  no  boss,  preparo- 
toid longitudinal  crest  present,  venter  often  spotted; 
middorsal  absent  or  present;  snout  6.1-9.1  in  L. 

American  toad,  Bufo  a.  americanus 
Plate  XXV 

J.  Brilliant  coloration;  long,  narrow  parotoids;  greater 
width  between  parallel  cranial  crests,  hind  limbs 
shorter,  ventral  granulation  smooth. 

Hudson  Bay  toad,  Bufo  americanus  cofei 
Plate  XXVI 

II.  Warts  round,  smaller,  parotoids  slightly  divergent  at 
rear  and  on  lateral  aspect,  parotoids  usually  in  contact 
with  postorbital  crest,  often  with  boss  on  nostril  with 
crests  extending  backward;  venter  unspotted  or  with 
median  pectoral  spot,  middorsal  line  present,  snout 
7.0-9.1  in  L. 

Rocky  Mountain  toad,  Bufo  w.  woodbousii 
Plate  XLIII 

CC.  Parotoids  round  or  triangular;  sole  (metatarsal)  tubercles  round,  small, 
noncutting;  dorsal  pattern  without  the  4-6  paired  spots  (of  the  JB. 
americanus,  w.  fowleri,  terrestns,  woodbousii,  etc.,  group). 
D.  Crest  prominent,  size  large,  £-84/5  in.  (50-1x0  mm.),  parotoid 
subtriangular. 

E.  Parotoid  as  large  or  larger  than  side  of  head,  divergent,  not  bi- 
colored;  toes  1/1-1/3  webbed,  crests  not  trenchant  and  top  of 
head  not  a  deep  valley,  brown  with  some  black,  yellow,  red, 
olive;  with  or  without  black  spots  on  a  light  vertebral  line; 
snout  7.3-8.6  in  L.  Marine  toad,  Bufo  marmus 

Plate  XXXVIII 


KEYS  87 

EE.  Parotoid  much  smaller,  not  as  large  as  side  of  head,  not  di- 
vergent, bicolored;  row  of  light  conical  tubercles  on  side,  body 
flat;  toes  1/3  webbed;  crest  high  trenchant  and  top  of  head  a 
deep  valley;  brown  or  blackish  brown  with  light  olive,  buff, 
or  cinnamon  area  down  back  and  a  similar  band  or  stripe  on 
either  side,  snout  6.6-10.4  in  L  Nebulous  toad,  Bufo  vallicefs 

Plate  XLII 

DD.  Crests  absent  or  obscure;  male  excrescences  on  first  two  fingers 
not  prominent,  size  small,  below  3  in.  (45  mm.). 
E.  Parotoids  large,  low,  descending  on  side,  as  long  as  side  of 
head,  body  rounded;  head  narrower  (1.7-3.15  in  L.);  snout 
distinctly  pointed  and  protruding  (6.9-9.1  in  L.),  foot  with 
tarsus  short  (1.0-1.4  m  L.),  size  small,  1-1  4/5  in.  (16-46 
mm.);  green  or  gray.  [Recently  subdivided  by  Taylor  into 
Girard's  two  forms,  B.  debths  and  B.  msidior.] 
(F.  Toes  about  1/3  webbed;  nostril  back  from  tip  of  snout; 
preorbital  and  suborbital  crests  smooth,  without  tuber- 
cles; parotoid  as  large  as  side  of  head;  anterior  edge  of 
tympanum  not  strongly  elevated. 

Little  green  toad,  Bufo  dcbtlts 
Plate  XXXIV 

FF.  Toes  about  1/1  webbed,  nostril  at  extreme  tip  of  snout; 
parotoid  larger  than  side  of  head;  anterior  edge  of  tympa- 
num elevated.  Bufo  msidtor 

Plate  XXXVII 
After  Smith  and  Taylor's  1948  checklist.) 

EE  Parotoids  small,  raised,  rounded,  body  flat,  head  broader  (1.3  - 
1.6  in  L.),  snout  not  distinctly  protruding,  canthus  rostrahs 
prominent,  foot  with  tarsus  medium  (1.66-1.88  in  L.),  size 
medium,  i  3/5-3  in.  (40-74  mm  );  red  to  gray. 

Spotted  toad,  Bufo  punctatus 
Plate  XXXIX 


Tree  Frogs,  HYLIDAE:  Acris,  Pseudacris,  Hyla 

A.  Alternating  dark  and  light  bands  on  rear  of  thigh;  oblique  white  stripe 
from  eye  to  shoulder;  vertical  dark  and  light  bars  on  upper  jaw;  white 
margined  triangle  between  eyes;  hind  leg  very  long  (0.55-0.61  in  L.); 
tibia  very  long  (1.5-1.7  in  L.).  Cricket  frog,  Acris  gryllus 


88 


HANDBOOK  OF  FROGS  AND  TOADS 


Figure  j.  Toads,  Bufonidae:  Bufo.  i.  Parotoid.  2.  Tympanum  (ear).  3.  Warts  at  angle 
of  mouth.  4.  Gland  on  femur.  5.  Glands  on  tibia.  6.  Two  metatarsal  (sole)  tubercles.  7. 
Two  metacarpal  (palmar)  tubercles.  8.  Crests  united  forming  a  prominent  raised  boss 
between  the  eyes.  9.  Canthus  rostralis.  10.  Canthal  crest,  n.  Preorbital  crest.  12.  Supra- 
orbital  crest.  13.  Postorbital  crest.  14.  Preparotoid  crest.  15.  Parietal  crest.  16.  Folds  of 
skin  of  lower  throat  of  male,  covered  at  periods  of  rest  by  the  lappet  (17).  17.  Lappet  or 
apron  at  rear  of  throat  of  male.  18.  Interparotoid  interval. 


Figure  4.  Tree  frogs,  Hylidae:  Acris,  Pseudacris,  and  Hyla.  i.  Tympanum  (ear).  2. 
Tympanic  fold.  3.  Snout  (muzzle).  4.  Plaits  on  throat  of  male.  5.  Pectoral  fold  across 
breast.  6.  Tarsal  fold.  7.  Small  adhesive  disks.  8.  Large  adhesive  disks.  9.  Prepollex.  10. 
Rear  of  casque  (skin  fastened  to  skull)  outlines  rear  of  head. 


KEYS  89 

(B.  Smaller;  less  web  (3  phalanges  of  toe  4  free,  toe  i  partly  free);  more 
rugose;  anal  warts  less  prominent;  legs  longer,  heel  reaching  beyond 
snout;  rear  of  thigh  more  definitely  striped.  Acris  g.  gryllus 

Plate  XLV 

BB.  Larger,  more  web  (i-i^  phalanges  of  toe  4  free,  toe  i  completely 
webbed);  smoother;  anal  warts  more  prominent;  legs  shorter,  heel 
not  reaching  to  snout;  rear  of  thigh  less  definitely  striped. 

Acris  g.  crepitans 
Plate  XLVI 
After  Dunn.) 

AA.  No  alternation  of  dark  and  light  bands  on  rear  of  thigh,  triangle  be- 
tween eyes,  if  present,  not  white-margined,  no  alternation  of  dark  and 
light  bars  on  upper  jaw. 

B.  No  dark,  brown,  black,  or  plum-colored  stripe  in  front  of  or  be- 
hind eye. 

C.  Head  skin  attached  to  skull;  thumb  rudiment  apparent;  rear  of 
femur  reticulated;  no  light  jaw  spot  or  dark  bar  between  eyes; 
disks  very  large,  size  large,  2.  3/5-5  1/5  in.  (64-130  mm.). 

Giant  tree  frog,  Hyla  stptentrionalis 
Plate  LXXI 

CC.  Head  skin  not  grown  to  skull;  no  thumb  rudiment;  rear  of  femur 
unspotted. 

D.  Rear  of  thigh  purple;  no  interorbital  bar  between  eyes; 
throat  with  green  on  either  side. 

E.  Green  of  female's  throat  not  edged  with  white;  back 

smooth  green;  body  slender  (head's  width  3.0-3.6  in  L.). 

F.  Yellow  or  white  line  along  side  and  on  upper  jaw; 

tibia  1.7-1.95  in  L. ,  third  finger  longer  (3.4-4.0  in  L.). 

Green  tree  frog,  Hyla  c.  cinerea 
Plate  LXIV 

FF.  No  yellow  or  white  line  along  side  or  on  upper 
jaw,  tibia  1.95-1.1  m  L.,  third  finger  shorter  (4.5- 
5.4  in  L.).  Miller's  tree  frog,  Hyla  c.  evittata 

Plate  LXV 

EE.  Green  of  female's  throat  edged  with  white;  back  usually 
granular  and  dark-spotted;  body  stout  (head's  width 
X.3-2..8  in  L.);  white  stripe  from  tip  of  snout  along  upper 
jaw  backward.  Barker,  Hyla  gratiosa 

Plate  LXIX 


90  HANDBOOK  OF  FROGS  AND  TOADS 

DD.  Rear  of  thigh  orange  or  ocher;  interocular  bar  present;  throat 
without  green  on  either  side. 

E.  Light  stripe  below  eye  to  shoulder,  back  black,  green,  or 
brown,  spotted  or  not;  smooth;  first  finger  7.0-9.3  in  L.; 
first  toe  7-14  in  L.;  interorbital  space  8.0-9.3  *n  L. 

Squirrel  tree  frog,  Hyla  squirella 
Plate  LXXII 

EE.  No  light  stripe  on  upper  jaw  or  light  spot  below  eye,  brown 
or  gray  usually  spotted;  large  disks;  first  finger  5.1-6.5  in  L.; 
first  toe  5.5-  8  8  in  L.;  interorbital  space  9.0-10.6  in  L. 

Canyon  tree  frog,  Hyla  arenicolor 
Plates  LX,  LXI 

BB.  Dark,  brown,  black  stripe  or  band  in  front  of  or  behind  eye  or  both. 
C.  Rear  of  thigh  spotted. 

D.  White-edged,  plum-colored  band  from  eye  to  groin;  back  green, 
unspotted  with  dark,  rear  of  femur  with  deep  orange  spots;  no 
interorbital  bar,  throat  with  green  on  either  side,  green  white- 
edged  in  female.  Anderson  tree  frog,  Hyla  andersonii 

Plate  LIX 

DD.  No  plum-colored  band;  interocular  bar  present,  back  usually 
with  spots. 

E.  Network  of  black  on  yellow  sides;  broad  dark  vitta  back 
from  eye  becoming  a  vertical  shoulder  bar,  throat  with 
greenish  or  yellow,  rear  of  thigh  netted  with  greenish  yellow 
and  purplish  russet;  light  yellow  or  green  spot  below  eye; 
size  large,  i  3/4-3  3/5  in.  (44-89  mm.). 

Mexican  tree  frog,  Hyla  baudinii 
Plate  LXIII 

EE.  No  network  of  black  on  yellow  sides;  no  black  or  brown 
shoulder  bar;  throat  not  prominently  greenish;  medium  to 
small. 
F.  Rear  of  thigh  brown,  no  netted  pattern. 

G.  Rear  of  thigh  with  distinct  round  or  elliptic  orange 
yellow  spots;  no  light  spot  below  eye,  a  cross-shaped 
spot  on  back.         Pine  woods  tree  frog,  Hylafemoralis 
Plate  LXVIII 

(GG.  Rear  of  thigh  specked  with  yellowish  brown  and 
darker  brown;  three  rows  of  approximated  spots  on 
back  or  cruciform  spot;  light  spot  below  eye  present 
or  absent.  (After  Cope.) 

Dusky  tree  toad,  Hyla  wrsicolor  pbaeocrypta} 

Plate  LXXVI 


KEYS  gt 

FF.  Rear  of  thigh  netted  with  black  or  dark;  light  spot  below  eye. 

G.  Rear  of  thigh  with  green  in  network;  groin  greenish; 

cross  on  back  in  center  or  rear  of  back;  back  less  rough; 

intertympanic  space  1.6-3.1  in  L.;  internasal  space  8-10 

in  L.;  third  finger  1.9-3.1  in  L. 

Viosca's  tree  frog,  Hyla  aviwca 
Plate  LXII 

GG.  Rear  of  thigh  with  orange  in  network;  groin  orange; 
cross  on  forward  half  of  the  back;  skin  commonly  rough; 
intertympanic  space  3.0-3.8  in  L.;  internasal  space  o.o- 
11.5  in  L.;  third  finger  3.1-4.8  in  L. 

Common  tree  toad,  Hyla  vcrsicolor 

H.  Dorsal  surfaces  smooth;   "a  number  of  subcircular 

golden  spots  in  the  brown  ground  on  rear  of  thighs; 

interspaces  [on  rear  of  femur]  often  reduced  to  small 

circular  spots."      Cope's  tree  frog,  Hyla  v.  chrysoscehs 

Plate  LXXIV 

HH.  Dorsal  surfaces  rough,  "brown  reticulation  on  yellow 
ground  of  the  posterior  face  of  the  thighs";  "more 
fully  marbled  with  yellow  and  brown,  even  covering 
the  whole  inner  face  of  the  tibia  and  the  light  inter- 
spaces more  or  less  angular." 

Common  tree  toad,  Hyla  v.  versicolor 

Plate  LXXIII 

CC.  Rear  of  thigh  unspotted,  usually  a  transverse  bar  or  triangle  or  median 
longitudinal  line  between  eyes. 

D.  A  narrow  oblique  cross  on  back;  rear  of  thigh  olive  ocher  or  raw 
sienna;  no  prominent  lateral  dark  stripe.  Peeper,  Hyla  crucifer 

(E.  Slight  and  indistinct  chest  spotting,  stripe  along  margin  of 
upper  jaw,  coloration  less  rich,  dorsal  stripes  less  broad. 

Peeper,  H.  c.  crucifer 
Plate  LXVI 

EE.  More  or  less  pronounced  ventral  spotting;  dark  spots  along 
margin  of  upper  jaw;  coloration  richer;  broader  dorsal  stripes. 

Florida  peeper,  H.  c.  bartramiana  Harper 
Plate  LXVII 

After  Harper.) 

DD.  No  oblique  cross  on  back,  usually  a  triangle  or  spot  between  eyes 
or  median  dark  longitudinal  line;  dorsal  and  vittal  stripes  or  rows 
of  spots. 


92  HANDBOOK  OF  FROGS  AND  TOADS 

E.  Size  very  tiny,  2.75-5/8  in.  (11.5-17.5  mm.);  three  dorsal  stripes  usually 
but  not  always  absent;  stripe  from  eye  backwards  usually  present;  very 
long  hind  limbs  (0.53-0.64  in  L.),  tibia  very  long  (1.5-1.1  in  L.). 

Little  chorus  frog,  Pseudacris  ocular  is 

Plate  LVI 

EE.  Size  small,  3/4-1  7/8  in.  (19-48  mm.);  4  or  5  dark  stripes  or  rows  of 
spots  usually  present;  hind  limb  0.61-0.87  in  L. 

F.  Five  stripes  or  rows  of  spots  usually  present,  size  small;  disks  in- 
conspicuous. 
Pseudacris  type  of  coloration  (middorsal,  dorsolateral,  and  vittal 

stripes  or  rows  of  spots,  5  in  number)  more  or  less  evident. 
A  dark  equilateral  interocular  triangle  (sides  concave  or  straight). 
Prominent  white  stripe  on  upper  jaw,  extending  backward  below 

prominent  vittal  stripe  and  more  or  less  along  side  of  body. 
A  dark  lateral  stripe  more  or  less  evident. 
No  oblique  cross  on  back  (occasionally  present  in  Pseudacris  brachy- 

phona  and  P.  streckerf). 
Tympanum  smaller,  usually  11-18  in  L. 
Winter  or  early  spring  breeders  (December-May). 
(G.  Lateral  black  stripe  sharply  defined  with  weak  dorsal  3-stripe 
pattern;  tendency  for  leg  markings  to  be  longitudinal;  narrow 
dark  line  on  outer  tibial  edge;  more  delicate,  smoother  skin; 
yellowish  venter.  (After  Brandt  and  Walker  )) 

Brimley's  chorus  frog,  Pseudacns  brimleyi 
Plate  XLVIII 

GG.  Each  lateral  stripe  and  3  dorsal  stripes  more  or  less  equally  de- 
fined; leg  markings  more  or  less  transverse,  small  size  (adults 
reaching  30,  31-37  mm.,  1.1-1.5  *n-)>  sexually  mature  at  19-11 
mm.  or  more,  Pseudacns  type  pattern  (5  stripes  or  rows  of  spots) 
generally  discernible;  middorsal  stripe  or  row  of  spots  usually 
present;  disks  small,  inconspicuous;  body  slender. 

Swamp  cricket  frog,  Pseudacns  nignta,  6  subspecies 

Plates  XLIX-LIV 

H.  Hind  limb  shorter  (0.74-0.81  in  L.,  av.  0.79);  tibia  shorter 
(1.36-1.8  in  L.,  av.  1.59),  foot  with  tarsus  shorter  (1.55-1.75 
in  L  ,  av.  1.69),  fore  limb  shorter  (i  06-1.11  in  L.,  av.  1.14); 
head  shorter  (head  to  tympanum  3.19-3.63  in  L.,  av.  3.40); 
mouth  shorter  (head  to  angle  of  mouth  3.73-5.71,  av.  4.39). 
Minn,  to  B.C.,  Great  Bear  Lake,  and  Hudson  Bay. 

Northern  striped  tree  frog,  P.  n.  septentrionalts 

Plate  LII 


KEYS  93 

HH.  Hind  limb  longer  (0.61-0.75,  avs.-o.64~o.7i),  tibia  longer  (1.87-1.54, 
avs.  1.94-1.36);  foot  with  tarsus  longer  (1.13-1.60  in  L.,  avs.  1.36- 
1.47);  fore  limb  longer  (1.5-1.33,  avs.  1.80-1.99),  head  longer  (head 
to  tympanum  1.55-3.19  in  L.,  avs.  i  94-3.14),  mouth  longer  (head  to 
angle  of  mouth  3.0-4.66,  avs.  3.39-3.71). 

I.  Tibia  shorter  (1.14-1.54  in  L.,  av.  1.36);  head  shorter  (3.14-3.19 
in  L.,  av.  3.14);  dorsum  finely  granular.  "All  the  upper  part  an 
inconspicuous  dark  greenish  gray  with  three  longitudinal  rows  of 
large  elongated  spots  .  .  ."  (Wied).  Middle  row  "unbroken  ...  to 
middle  of  the  back,"  then  "separate  spots. ff  N.Y.  and  Ont.  to 
Idaho  and  northern  Ariz.  Striped  tree  frog,  Pscudacris  n.  tnsmata 

Plate  LIII 

II.  Tibia  longer  (1.77-1.16  in  L),  head  longer  (1.55-3.19  in  L.,  av. 

i  94-3-04)- 

J.  Head  shorter  (av.  3.04  in  L.),  head  wider  (av.  3.13  in  L.).  "Above 
dark  or  fawn,  with  three  nearly  parallel  stripes  down  the  back, 
the  central  widening  but  scarcely  bifurcate  behind  .  .  ."  (Baird). 
Dorsum  more  or  less  smooth.  N.J  to  N.C.,  possibly  S.C. 

Eastern  chorus  frog,  Pseudacris  n.  fcriarum 
Plate  LI 

JJ.  Head  longer  (av.  1.94  in  L.);  head  narrower  (av.  3.18-3.6  in  L.). 

K.  Snout  shorter  (5.71-70  in  L,  av.  633).   "Above  grayish 

brown  or  ash  with  distinct  large  circular  blotches"  (Baird). 

Dorsum  smooth  or  only  slightly  tubercular.  Texas  to  Kansas. 

Clarke's  chorus  frog,  Pseudacns  n.  clarkii 

Plate  L 

KK.  Snout  longer  (4.6-6.66  in  L.,  av.  5.70),  dorsum  more  or  less 
tubercular. 

L.  "A   light   line   is   present   along    the    jaw."    "Above, 

speckled  with  small  white  warts,  middle  of  back  cinereous, 

with  an  interrupted  stripe  of  black"  (Le  Conte).  N.C.  to 

La.  Swamp  cricket  frog,  Pseudacns  n.  nigrita 

Plate  XLIX 

LL.  "Upper  lip  dark  plumbeous  with  a  series  of  ...  white 
spots"  (Cope)  or  these  absent.  (Dorsum  often  tubercular, 
hind  limb  light  intervals  broader  than  in  P.  n.  ni&nta. 
Dorsal  spots  smaller  and  better  separated — Brady  and 
Harper).  Florida. 

Florida  chorus  frog,  Pseudacris  n.  wrrucosa 
Plate  LIV 


94  HANDBOOK  OF  FROGS  AND  TOADS 

FF.  Four  stripes  or  rows  of  spots. 

G.  Vittal  stripe  slender,  dorsolateral  bands  curved,  sometimes 
making  a  cross  or  transverse  bar  on  back;  hind  limbs  long 
(0.60^-0.67  in  L.),  triangle  between  eyes;  disks  distinct. 

Mountain  chorus  frog,  Pseudacris  bracbyphona 
Plate  XLVII 

GG.  Vittal  mask  white-bordered  above;  vittal  stripe  darker  than 
other  stripes  or  rows  of  spots;  hind  limb  0.61-0.87  in  L. 
H.  Body  broad,  toadlike  (1.75-1  o  in  L.);  disks  inconspicuous; 
vitta  ends  at  shoulder,  hind  limbs  shorter  (o  73-  0.87  in  L.). 
Strecker's  ornate  chorus  frog,  Pseudacris  streckeri 
Plate  LVIII 

HH.  Body  narrow  (1.94  in  L.,),  hind  limbs  longer  (o  61-0.78 
in  L.). 

I.  Disks  inconspicuous,  dark  mask  ending  beyond  shoulder 
sometimes  to  groin,  oblique  groin  spots  light-bordered, 
interorbital  space  narrow  (8.8-14.4  in  L.). 

Ornate  chorus  frog,  Pseudacris  ornata 
Plate  LVII 

II.  Disks  distinct,  interorbital  wide  (7.1-10  in  L  ),  no  light 
borders  around  groin  spots  if  present. 
J,  Vitta  may  extend  some  distance  along  the  side,  inter- 
ocular  triangle  or  bar  absent,  dark  spot  each  eyelid 
usually  present,  a  pair  of  dark  longitudinal  postsacral 
bars  or  spots — the  conspicuous  dorsal  marking,  tym- 
panum smaller  (14.4-10  in  L.),  first  finger  shorter 
(7.10-10  in  L.),  first  toe  shorter  (7.0-13.3  in  L.), 
anterior  edge  of  tibia  with  heavy  brown  spots,  lack- 
ing white  line.      Wrights'  tree  frog,  Hyla  wnghtorum 

Plate  LXXV 

JJ.  Vitta  commonly  ends  at  shoulder;  interocular  triangle 
usually  present,  no  very  distinct  postsacral  bars,  the 
dorsal  spotting  often  quite  pronounced  and  varied, 
tympanum  larger  (11-14  *n  L.),  first  finger  longer 
(5.6-8.0  in  L.),  first  toe  longer  (5.6-8.8  in  L.). 

Pacific  tree  frog,  Hyla  regilla 
Plate  LXX 

Robber  Frogs,  LEPTODACTYLIDAE:  Leptodactylus, 
Eleutherodactylus,  Syrrhophus 

A.  Pupil  horizontal,  tympanum  distinct. 

B.  Fingers  and  toes  free  without  distinct  terminal  disks,  toes  without 


KEYS 


95 


dermal  border,  size  small,  i  1/5-1  in.  (15-49  mm.);  a  lateral  fold; 
white  band  along  jaw  to  canthus  oris  and  humerus;  dorsal  color  choco- 
late-brown, limbs  dark  cross-barred,  two  small  tarsal  tubercles;  narrow 
tarsal  fold,  heel  to  orbit.  White-lipped  frog,  Leptodactylus  labtalis 

Plate  LXXXII 

BB.  Tips  of  phalanges  T-shaped,  toes  and  fingers  free;  terminal  disks  small, 
no  white  upper  jaw  stripe,  no  dorsolateral  or  lateral  fold. 
C.  Size  large,  i  7/8-3  3/5  in.  (48-90  mm.),  a  ventral  disk,  voice  a  re- 
sounding bark,  head  broad  (1-1.57  in  L.),  head  wider  than  head  to 
tympanum;  eye  small  (7.1-9.15  in  L.),  eye  much  less  than  first  finger; 
fingers  larger,  light  stripe  down  middle  of  back,  intertympanic  fold 
present  (in  preserved  material). 

D.  Fourth  toe  longer;  fairly  closely  aggregated  black  blotches,  sides 
and  hinder  half  of  abdomen  faintly  areolate,  skin  in  adults  on 
dorsum  stiff,  coarse,  areolate.  Belly  without  vermiculations  or 
spots,  back  covered  with  large  black  spots. 

Texas  cliff  frog,  Eleuthcrodactylut  latrans 
Plates  LXXIX-LXXX 

[DD.  Fourth  toe  shorter,  upper  parts  either  green  or  with  scattered 
tubercles,  belly  with  faint  spots,  broad  light  transverse  band 
across  back  just  back  of  fore  limbs. 

Mexican  cliff  frog,  Elcutherodactylus  augusti 
Plate  LXXVII] 

CC.  Size  small,  3/5  i  3/5  in.  (15-39  5  mm),  without  ventral  disk,  voice 
a  cnckethke  chirp;  head  narrower  (i  66-3.38  in  L.),  head  usually 
narrower  than  head  to  tympanum,  eye  larger  (5-8  in  L.),  eye  greater 
than  first  finger,  fingers  smaller. 

D.  Usually  a  light  stripe  from  eye  backward  along  dorsolateral  re- 
gion, a  light  transverse  band  between  eyes,  toes  longer,  fore  limb 
shorter  (i  7-1.0  in  L.),  foot  with  tarsus  longer. 

Ricord's  frog,  Elcutherodactylus  rtcordtt  plantrostns 
Plate  LXXVIII 

DD.  Usually  without  light  dorsolateral  stripe  from  eye  backward, 
usually  no  transverse  band  between  eyes;  toes  shorter;  fore  limb 
longer  (i  45-1.88  in  L.). 

E.  Tympanum  smaller  (11-13  *n  *-.)>  hind  limb  shorter  (0.71  - 
0.87  in  L.),  internasal  broader  (8-9  in  L.);  fore  limb  usually 
greater  than  foot  with  tarsus. 

Marnock's  frog,  Syrrhopbus  marnockti 
Plate  LXXXIV 

F.  Vermiculatc  dorsal  pattern,  wider  head  (ratio  to  L.  0.40), 
smaller  size  (11-18  mm.)  Chisos  Mts. 

Gaige's  frog,  Syrrhophus  gaigeae 


96  HANDBOOK  OF  FROGS  AND  TOADS 

FF.  Spotted  dorsal  pattern,  narrower  head  (ratio  of  width 
of  head  in  L.  0.36);  larger  size,  2.5-35  mm-  Central  Texas. 
Marnock's  frog,  Syrrhopbus  mamockii 
Plate  LXXXIV 

EE.  Tympanum  larger  (8-10  in  L.),  hind  limb  longer  (o-72.-o.73 
in  L.),  internasal  narrower  (9-10  in  L  ),  fore  limb  usually 
less  than  (rarely  equal  to)  foot  with  tarsus. 

Camp's  frog,  Syrrhophus  campi 
Plate  LXXXIII 


Frogs,  RANIDAE:  Rana 

L.  Tympanum  larger  than  eye  in  male,  equal  in  female  (rarely  smaller  in 
R.  wr&attpcs),  5 .6-11.5  *n  L. ,  throat  of  male  differently  colored,  upper  jaw 
unicolor,  no  black  mask,  no  light  jaw  stripe,  no  regular  dorsal  rows  of 
spots,  no  pronounced  yellow,  orange,  or  red  on  undersides  of  hind  limbs 
or  on  belly  or  groin,  tibia  i  8  2.  6  in  L. 

B.  Rear  of  femur  with  alternations  of  light  and  dark  horizontal  bands; 
dorsolatcral  folds  absent,  edge  of  jaw  uniform. 

C.  Size  small  i  5/8-1  5/8  in  (41  66mm  ),  usually  a  light  yellowish 
dorsolatcral  stripe  extends  back  from  eye,  two  joints  of  fourth  toe 
free  of  web,  vocal  sacs  of  male  like  a  marble  on  either  side  of  head, 
intcrtympanic  space  3. 41-  4. 4  in  L.,  foot  with  tarsus  i  36-1. 56  in  L. 

Sphagnum  frog,  Rana  vtrgattpes 
Plate  CXXIII 

CC.  Size  Urge  3  1/4  8  in.  (82.  2.00  mm  ),  no  or  only  one  joint  of  fourth 
toe  free  of  web,  back  uniform  in  color,  no  vocal  sacs  in  males  on 
side  of  head 

D.  Alternation  of  color  on  rear  of  femur  conspicuous,  first  linger 
generally  less  than  second  finger,  mtertympanic  space  narrower 
(50  6  8  in  L  ),  foot  with  tarsus  shorter  (i  4-1  67  in  L  ). 

Southern  bullfrog,  Rana  grylto 
Plate  XCVII 

DD  Alternation  of  color  on  rear  of  femur  not  conspicuous,  first 
finger  generally  equal  to  second,  mtertympanic  space  wider 
(4.5-5.1  in  L  ),  foot  with  tarsus  greater  (1.18-1.53  in  L  ). 

Bullfrog,  Rana  catcsbctana 
Plate  XCIV 

BB.  Rear  of  femur  without  alternation  of  light  and  dark  horizontal  bands; 
edge  of  jaw  mottled,  barred,  or  uniform. 

C.  Dorsolateral  fold  absent,  size  large,  31r-5J4  in.  (81-131  mm); 
rear  of  femur  light  or  white  spots  on  brown,  edge  of  upper  jaw 


KEYS 


97 


5- 


Figttre  5.  Robber  frogs,  Leptodactylidae:  l^ptodactylus,  Elcuthetodactylus,  and  Syr- 
rophus.  i.  Ventral  disk.  2.  Dorsolatcral  fold.  3.  Lateral  folds.  4.  Transverse  (T-shaped) 
disks.  5.  Subarticular  tubercles  sharp  and  saw-toothed.  6.  Brachium  (upper  arm).  7. 
Antebrachium  (forearm).  8.  Femur.  9.  Tibia.  10.  Heel.  n.  Tarsus.  12.  Foot. 


10 


*Figute  6.  Frogs,  Ranidae:  Rana.  T.  Nostril.  2.  Internasal  space.  3.  Costal  (dorsolateral) 
fold.  4.  Enlarged  thumb  of  male.  5.  Enlarged  tympanum  of  male.  6.  Tympanum  of  fe- 
male 7.  Tympanic  fold.  8.  Fleshy  fold  on  jaw.  9  Glandular  folds  on  tibia.  10.  Sacral 
hump.  IT.  Full  webbing  of  male  bullfrog.  12.  Narrow  mtcrorbital  space.  13.  Intertympanic 
space. 


Ft gut e  7.  Narrow-mouthed  toads,  Brevicipitidae:  Hypopachus  and  Microhyla.  i.  Broad 
waist.  2.  Femur  partly  involved  in  body  skin.  3.  Two  metatarsal  tubercles.  4.  Slight  basal 
web.  5.  Transverse  fold  of  skin  across  head.  6.  Depressed  form  of  body.  7.  Body  thick. 
8.  Legs  short. 


98  HANDBOOK  OF  FROGS  AND  TOADS 

mottled;  head  to  angle  of  mouth  1.0-1.7  in  L.;  first  finger 
6.5-7.3  in  L.  River-swamp  frog,  Rana  hcckscheri 

Plate  XCVIII 

CC.  Dorsolateral  fold  present,  indistinct,  or  absent;  edge  of  upper 
jaw  uniform  or  barred;  size  medium  to  small;  first  finger  5.0-6.8 
in  L.;  head  to  angle  of  mouth  1.7-3. 4  *n  L. 
D.  Dorsal  folds  lacking  or  interrupted;  back  and  sides  mottled 
or  with  prominent  spots,  rear  of  femur  vermiculated  in  its 
spotting;  males  with  lateral  external  vocal  sacs  somewhat 
developed,  size  small,  i  7/8-3  in.  (48-76  mm.),  third  toe 
1.5-1.9  in  L.;  interorbital  space  19-18  in  L. 

Mink  frog,  Rana  septtntrionalis 
Plate  CXVIII 

DD.  Dorsal  fold  on  cephalic  half  only,  cheek  green  with  mottled 
jaw  below;  back  uniform  or  with  fine  black  speckings;  rear 
of  femur  with  fine  and  scant  speckings;  males  with  no  lateral 
external  vocal  sacs;  size  medium,  1-4  in.  (51-100  mm.);  third 
toe  1.7-3.7  in  L.;  interorbital  space  11-13  *n  L. 

Green  frog,  Rana  clamitans 
Plate  CXV 

AA.  Tympanum  of  male  not  enlarged  (enlarged  in  JR.  onca,  rarely  in  R. 
artolata  and  R.  sphtnocepbala),  smaller  than  eye  (9-19  in  L.),  throat  not 
so  differently  colored  in  males;  upper  jaw  with  light  stripe  or  mottling 
of  light  and  dark,  a  vitta  or  regular  spots  between  dorsolateral  folds  or 
with  yellow,  orange,  or  red  on  hind  limbs,  groin,  or  belly,  tibia  1.4- 
1.3  in  L. 

B.  Dorsolateral  folds  absent  or  indistinct,  low,  broken;  under  surface 

of  hind  limbs  yellow;  skin  rough,  no  vitta,  body  odor  pronounced. 

C.  Size  larger,  1 1/3-4  1/1  in.  (58-115  mm.);  no  outer  sole  (meta- 

tarsal)  tubercle;  no  stripe  on  upper  jaw;  throat  and  lower  jaw 

uniform  or  cloudy,  hind  limbs  shorter  (0.65-0  69  in  L.),  tibia 

shorter  (1.86-1.0  in  L.).  Mexican  frog,  Rana  tarabumarae 

Plate  CXXII 

CC.  Size  smaller,  i  3/5-3  3/8  in.  (3 9-84 mm.);  outer  sole  (metatarsal) 
tubercle,  stripe  on  upper  jaw  present  or  obscure,  throat  and 
lower  jaw  spotted,  hind  limbs  longer  (0.57-0.66  in  L.);  tibia 
longer  (1.61-1.88  in  L.)  Yellow-legged  frog,  Rana  boylit 

Plates  LXXXIX-XCI 

("Vomerine  teeth  rudimentary,  on  two  oblique  ridges  between 
the  nares  .  .  .  ;  tympanic  region  not  darker  than  rest  of  head; 


KEYS 


99 


fold  along  upper  lip  colored  like  rest  of  body,  mottled  or  dark; 
red  never  present  in  coloration.  Rana  boylti  and  subspecies" 
"i.  When  hind  leg  is  brought  forward  along  the  body,  inside  angle 
of  bent  tarsus  and  tibia  reaches  at  least  to  nares  and  often  beyond  the 
end  of  the  snout,  tympanum  covered  with  many  hispid  points. 
"3.  A  light  patch  on  top  of  head,  darker  area  crossing  the  posterior 
half  of  each  upper  eyelid  merging  insensibly  into  dorsal  color  be- 
hind; body-length  under  70  millimeters. 

[California  yellow-legged  frog,]  Rana  boyln  boylti  Baird" 
Plate  LXXXIX 

"3F.  No  light  patch  on  top  of  head,  darker  areas  crossing  posterior 
half  of  each  upper  eyelid,  when  present,  contrasting  with  dorsal 
coloration,  body  length  reaching  81  millimeters. 

[Sierra  Madrc  yellow-legged  frog,]  Rana  boyln  mutcosa  Camp" 
Plate  XC 

"2..  When  leg  is  brought  forward,  inside  angle  of  bent  tarsus  seldom 
reaching  beyond  nares,  tympanum  smooth  or  with  but  a  few  hispid 
points;  no  light  patch  on  top  of  head;  body  length  reaching  73 
millimeters. 

[Sierra  Nevada  yellow-legged  frog,]  Rana  boylti  sierrae  Camp" 
Plate  XCI 

C.  L  Camp,  Calif.,  1917,  p.  113.) 

BB.  Dorsolatcral  folds  distinct  full  length  of  body  (folds  not  conspicuous  in 
R.  aurora?)^  usually  with  a  mask;  skin  smoother,  body  odor  not  pro- 
nounced. 

C.  With  mask  to  angle  of  jaw  (sometimes  less  conspicuous  or  absent  m 
R.  aurora);  males  with  no  external  vocal  sacs  between  ear  and  shoul- 
der, rear  of  femur  finely  dotted  and  without  heavy  spots. 

D.  No  red  or  yellow  on  the  under  parts,  mask  black  or  dark  brown, 
size  small  i  1/6-3  */4  *n-  (19^82.  mm.),  tympanum  larger  (11- 
15  in  L.) 

E.  Dorsal  color  usually  a  mid-band  of  darker  color  within  light 
dorsolateral  folds,  middorsal  light  stripe  present  or  absent, 
breast  more  spotted.  Hind  limb  short  (0.62.^0  75  in  L.),  tibia 
short  (i.93'-2..3  in  L.);  tibia  usually  equals  foot. 

Northern  wood  frog,  Rana  sylvatica  cant abri gens ts 

Plates  CXX,  CXXI 

EE.  Dorsal  color  between  dorsolateral  fold  usually  like  dorsum; 
no  middorsal  stripe;  breast  usually  without  spots;  hind  limb 
long  (0.53-0.62.  in  L.);  tibia  long  (1.6-1.88  in  L.);  tibia 
longer  than  foot.  Wood  frog,  Rana  s>  sylvatica 

Plate  CXIX 


ioo  HANDBOOK  OF  FROGS  AND  TOADS 

DD.  Some  red  or  yellow  on  under  parts;  size  average  larger,  i  4/5-5  in.  (45- 

115  mm.);  tympanum  larger  (11-17  *n  L.)  or  smaller  (17-11  in  L.). 

E.  Angle  of  tarsus  and  tibia  reaching  to  eye,  not  beyond  nostril;  no 

mottling  in  groin,  mask  absent  (if  present  obscure  brown),  inky 

spots  on  back  sometimes  light-centered,  no  prominent  white  strip 

from  snout  or  eye  to  shoulder;  tympanum  small  (16.4-11  in  L.); 

eggs  with  i  jelly  envelopes,  mternasal  space  shorter  (11-16.4  *n  L.)- 

F.  Two  metatarsal  tubercles,  palmar  tubercles  present;  back  and  top 

of  head  with  inky  black  spots,  tympanum  smaller  (16.4-11  in 

L.);  interorbital  space  larger  (11-17.6  times  in  L.),  fingers  longer, 

especially  first  (6.1-6.45  in  L.)  and  second  (6.0-6.45  in  L.),  upper 

eyelid  narrower  (14.4-11  in  L.). 

Western  spotted  frog,  Rana  p.  prcttosa 
Plate  CXVI 

FF.  One  metatarsal  tubercle,  the  outer  absent,  palmar  tubercles  small 
or  wanting;  few  irregular  dorsal  spots,  sometimes  considerable 
spotting;  under  parts  yellow  or  deep  orange,  tympanum  larger 
(13-16  times  in  L.);  interorbital  space  smaller  (16.1-19  i  in  L.); 
finger  shorter,  especially  first  (5  8-7.8  in  L.)  and  second  (5  8-7.8 
in  L.),  upper  eyelid  wider  (11.6-16  in  L.). 

Nevada  spotted  frog,  Rana  p.  luteiventris 
Plate  CXVII 

EE.  Angle  of  tarsus  and  tibia  reaching  nares  or  beyond  end  of  snout;  al- 
ways or  slightly  mottled  in  groin,  mask  prominent  black  or  dark 
brown  (rarely,  absent),  prominent  stripe  from  eye  or  snout  to 
shoulder  or  shorter,  tympanum  normally  larger  (11-17  ln  L.),  eggs 
with  3  distinct  jelly  envelopes,  internasal  space  greater  (8.6  14.1 
in  L.). 

F.  Slight  mottling  in  groin.  Dorsum  greenish  brown  with  many 
black  spots;  under  parts  light  yellow  or  pale  salmon  orange; 
third  finger  shorter  (4  08-5.17  in  L.);  hind  limb  slightly  shorter 
(0.61-0.73  in  L.),  fourth  toe  slightly  shorter  (1.9-1.45  m  L.). 
Cascades  in  Washington  and  northern  Oregon  4000  ft.  or  higher. 

Slater's  frog,  Rana  cascadac 
Plate  XCIII 

FF.  Prominent  mottling  in  groin;  dorsum  without  inky  or  black 
spots  (some  few  light-centered)  under  parts  red,  third  finger 
longer  (3.16-4.71  in  L.),  hind  limb  slightly  larger  (0.50-0.69  in 
L.);  fourth  toe  slightly  longer  (1.51-1.15  m  L.). 

Rana  aurora  group 
Plates  LXXXVII,  LXXXVIII 

G.  Skin  smooth;  dorsolateral  folds  distinct;  dorsum  without 
spots  or  dotted,  head  to  tympanum  in  width  of  head  0.93- 


KEYS  ioi 

1.03,  head  narrower  (z  5-3 .z  in  L.),  intcrtympanic  space 
(3.5-5.1  in  L.);  tibia  shorter  (1.67-1.53  in  L.),  first  toe 
(5.4-8.0  in  L.)  and  second  toe  (3.4-4.85  in  L.)  shorter; 
size  smaller,  i  3/4-3  i/z  in.  (44-87  inches). 

Oregon  red-legged  frog,  Rana  a.  aurora 

Plate  LXXXVII 

GG.  Skin  rough,  dorsolateral  folds  distinct;  dorsum  with  light- 
centered  spots,  head  to  tympanum  in  width  of  head  0.98- 
1.2.5,  s*2e  larger,  z 1/1-5  L/5  m-  (5 8-136  mm.);  head  wider 
(z  15-3  o  in  L.),  intertympanic  space  wider  (3.0-4.45  in 
L.),  tibia  longer  (i.46-zo6  in  L.),  first  toe  (4.5-6.5  in 
L  )  and  second  toe  (z  9-4.4  in  L  )  longer.  California  and 
Lower  California.  Introduced  into  Nevada. 

California  red-legged  frog,  Rana  a.  draytomi 

Plate  LXXXVIII 

CC.  With  no  mask,  males  with  external  vocal  sacs  between  car  and  shoulder. 
D.  Without  white  line  on  upper  jaw. 

E.  Upper  jaw  uniform  or  with  few  dashes,  size  small,  i  3/4-3  in. 
(44-74  mm  ),  dorsum  unspotted  or  with  more  numerous  spots 
than  R.  piptcns;  with  yellow  on  under  parts;  males  with  enlarged 
tympana,  hind  limbs  shorter  than  in  Rana  onca  (1.1-1.5  i*1  L.)- 

Nevada  frog,  Rana  fishen 
Plates  XCVI,  CVIII 

F.  "Fewer  dorsal  spots  and  much  shorter  hind  legs"  (Linsdale). 
Hind  limbs  longer  than  R.  fishert  (1.46-1.8  in  L.),  no  enlarged 
tympana.  Utah  frog,  Rana  onca 

Plates  XCIX,  CIX 

[The  Rana  fishert  of  Las  Vegas  region,  Rana  onca  of  Utah  and 
Nevada,  and  Rana  ptptens  of  Utah,  Nevada,  Arizona,  and 
Imperial  Valley  need  close  examination.] 

EE.  Upper  jaw  mottled,  size  larger,  zJ^S^  in.  (63- -113  mm.),  three 
or  four  rows  of  spots  between  dorsolateral  folds;  males  rarely 
with  enlarged  tympana. 

F.  Dorsal  spots  large,  light-centered,  round  with  light  borders, 
throat  and  chin  clear  except  at  sides;  three  or  four  crossbars 
on  legs  with  small  intermediates;  snout  shorter,  head  to 
tympanum  z.8  to  3.z  in  L.;  head  to  angle  of  mouth  3.15-3.8 
in  L.;  eye  small  (9.8-^.3  in  L.),  intertympanic  space  medium 
(3.8-4.7  in  L.),  width  of  head  in  L.  narrower  (z.6  to  z.gz  in 
L). 

(G.   "Head  U-shaped  in  outline  when  viewed  from  above; 
dorsum  often  smooth,  or  nearly  so;  tibia  length  less  than 


102  HANDBOOK  OF  FROGS  AND  TOADS 

40  mm.  in  adults,  posttympanic  fold  poorly  developed,  dorso- 
lateral  folds  narrow  or  only  slightly  raised  or  both. 

[Texas  gopher  frog,]  liana  areolata  areolata  Baird" 

Plate  LXXXV 

GG.  "Head  orbiculate  in  outline  when  viewed  from  above;  dorsum 
rugose,  tibia  length  more  than  40  mm.  in  adults;  posttym- 
panic fold  well  developed;  dorsolateral  folds  prominent. 

[Northern  gopher  frog,]  Rana  areolata  ctrculosa 

Rice  and  Davis" 

Plate  LXXXVI 
After  Goin  and  Netting,  La.,  1940,  p.  146.) 

FF.  Dorsal  spots  small,  irregular  slightly  or  not  light-encircled;  5  or 
8  uniform  bars  on  hind  limbs,  snout  longer,  head  to  tympanum 
i. 2.^3.0  in  L.,  head  to  angle  of  mouth  2-5-3  *  in  L->  eye  large 
(6.8-10.3  in  L.),  intertympanic  space  broad  (^.95-3. 65  in  L.); 
width  of  head  in  L.  wider  (i  93  -  2.. 6  in  L.)>  venter  of  head  heavily 
spotted. 

(G.  "Head  triangular  in  outline,  dorsolateral  folds  high  and  rela- 
tively narrow,  dorsum  with  numerous  prominent  warts; 
dorsal  spots  poorly  differentiated  from  gray,  brown,  or  black 
ground  color,  venter  always  spotted  at  least  from  chin  to 
midbody.,  dark  bars  on  hind  limbs  separated  by  interspaces 
that  are  never  wider  than  the  bars. 

[Dusky  gopher  frog,]  Kana  sevosa  (Goin  and  Netting) 

GG.  "Head  subtriangular  in  outline,  dorsolateral  folds  low  and 
very  broad;  dorsum  smooth  or  lightly  warty;  dorsal  spots 
distinct  against  pale  ground  color,  chin  and  throat  spotted; 
belly  usually  immaculate  posteriorly,  dark  bars  on  hind 
limbs  separated  by  light  interspaces  that  are  wider  than  the 
bars."  Gopher  frog,  Ratia  capito 

Plate  XCII 
After  Goin  and  Netting,  La.,  1940,  p.  146.) 

DD.  With  white  line  on  upper  jaw. 

E.  No  orange  on  under  parts,  dorsal  spots  round  with  interspaces  of 
same  diameter  or  more. 

F.  Snout  shorter  (60-6.8  in  L),  more  lateral  spots  below  dorso- 
lateral fold;  tibia  shorter  (1.73-1.94  in  L.);  head  to  tympanum 
z.8-3.z  in  L.;  upper  eyelid  wide  (n.i-i4  in  L.),  tympanum 
normally  without  light  center. 

Meadow  frog,  Kana  p.  pipiens 

Plate  CI 

[R.  pipiens  complex.  Plates  CV-CXV.] 


KEYS 


103 


FF.  Snout  longer  (5 .2.3-6.3  in  L.);  fewer  lateral  spots  below 
dorsolatcral  fold;  tibia  longer  (1.55-1.82.  in  L.),  head 
to  tympanum  2..3&-2..S  in  L. ,  upper  eyelid  medium  (9.3- 
11.7  in  L.),  tympanum  usually  with  light  center. 

Southern  meadow  frog,  Rava  p.  sphenoccphala 
Plate  CIV 

EE.  Orange  on  under  parts;  dorsal  square  spots  in  regular  rows; 
spots  with  interspaces  less  than  diameter  of  spots. 

Pickerel  frog,  Raua  palustris 
Plate  C 

Narrow-mouthed  Toads,  BREVICIPITIDAE:  Hypopachus, 

Microhyla 

A.  Two  sole  (metatarsal)  tubercles;  basal  webs  on  feet;  snout  shorter 
(9-10  in  L.);  usually  a  middorsal  yellow  line  and  mid-ventral  white 
line,  oblique  white  band  from  eye  to  shoulder. 

Taylor's  toad,  Hypopacbus  cumus 
Plate  CXXIV 

AA.  One  sole  (metatarsal)  tubercle,  no  webs  on  feet,  snout  longer  (6.6  8.2. 
in  L.),  no  middorsal  or  mid-ventral  lines  nor  oblique  postorbital  band. 
B.  Muzzle  shorter;  hind  foot  unusually  short,  hind  limb  short,  back 
areolate,  the  posterior  parts  even  pustular. 

Mitchell's  narrow-mouthed  toad,  Mtcrobyla  artolata 
BB.  Muzzle  longer,  hind  limb  longer. 

C.  Body  depressed  (thickness  in  length  3.0-5.6  in  males,  4.0  -4.3  in 
females),  tibial  width  in  tibial  length  3-4,  body  width  in  females 
2.  46-  3.1  in  L.;  upper  eyelid  10-18  in  L.,  eye  8.7-1^.7  in  L.;  skin 
usually  smooth,  under  parts  white,  dorsum  grayish  olive. 

Texas  narrow-mouthed  toad,  Mtcrohyla  oltvacea 
Plate  CXXVI 
(D.  Ventrum  immaculate  or  with  scattered  melanophores. 

E.  A  blotch  or  spots  on  the  femur  and  tibia  which  form  a  bar 
or  continuous  line  when  the  limb  is  folded,  dorsum  with 
dark  spots.  Taylor's  Microhyla  ma^atlanemts 

EE.  Spots  rarely  present  on  the  femur  and  tibia,  if  present  usu- 
ally not  forming  a  distinct  bar  when  the  limb  is  folded, 
dorsum  tan  and  generally  without  markings.        olivacea. 
After  M.  K.  Hecht  and  B.  L.  Matalas,  Gen.,  1946,  p.  7.) 
CC.  Body  less  depressed  (2-5-3.35  in  males,  z. 65-3. 4  in  females), 
tibial  width  in  tibial  length  2-5-3  -°  times,  body  width  in  females 
1.83-1.3  in  L.,  upper  eyelid  i8.6-z8  m  L.,  eye  icx-i4  m  L.,  skin 
smooth,  tuberculate,  or  pustular;  under  parts  gray,  or  brown 
speckled  and  mottled;  dorsum  black,  gray,  or  brown. 

Narrow-mouthed  toad,  Microhyla  carolinensis 
Plate  CXXV 


Accounts  of  Species 

ORDER  SALIENTIA  Laurenti 
FAMILY  ASCAPHIDAE 

Genus  ASCAPHUS  Stcjneger 

Map  4 


THE  first  recognition  of  this  form  by  Dr.  Stejneger  and  the  subsequent 
studies  of  it  by  Dr.  Helen  Thompson  Gaige  were  two  of  the  outstanding 
events  in  North  American  batrachology  in  the  last  forty-five  years.  This 
"tailed"  frog  has  relatively  the  broadest  internasal  space  of  all  our  frogs,  4.66 
to  8  into  length,  whereas  all  others  range  from  8  (or  7.77)  to  21.4  into  length.  In 
hind  limb  it  is  relatively  longer  than  robber  frogs,  narrow-mouths,  spadefoots, 
or  toads  and  comparable  to  tree  frogs  and  frogs.  It  has  long  fore  limbs.  Pos- 
sibly both  fore  and  hind  limbs  are  long  for  climbing.  Of  the  first  live  ones  we 
had,  all  except  one  escaped  by  crawling  up  the  inner  sides  of  a  barrel.  It  has 
a  large  mouth,  snout,  eye,  interorbital  space,  and  upper  eyelid. 

The  relationships  of  this  frog  have  been  much  discussed.  When  Stejneger 
(1899)  with  rare  courage  and  fine  discrimination  described  this  form,  he 
called  it  a  Discoglossid.  Later,  VanDenburgh  (1912)  found  it  to  be  not  "ecau- 
date," and  followed  Stejneger.  American  authors  such  as  Barbour  and  Coch- 
ran  (1932)  and  most  foreign  workers  like  Boulenger  (1910),  Gadow  (1920), 
Nieden  (1923),  and  Perrier  (1925)  kept  it  within  the  Discoglossidae.  From 
1924  to  1931  and  until  his  death  Noble  placed  it  in  the  Liopelmidae,  and  Vil- 
liers  (1934)  and  Pusey  (1943)  are  in  agreement  with  this  view.  In  1923  Fejer- 
vary  created  a  special  family,  the  Ascaphidae,  and  in  the  third  to  fifth  editions 
of  Stejneger  and  Harbour's  check  list  (1933,  1939,  and  1943)  these  authors 
adopt  this  family  name,  which  we  now  follow.  We  have  seen  considerable 
material  both  preserved  and  alive  and  are  much  impressed  with  its  Discoglos- 
sid affinities. 


RIBBED  TOADS:  ASCAPHIDAE  105 

American  Bell  Toad,  American  Ribbed  Toad 

Ascaphus  truet  Stejneger.  Plate  XVI;  Map  4. 

Range:  Washington,  Oregon,  northern  California  (south  to  Dyerville; 
Myers,  1931,  1943),  Emida,  Idaho  (Ehrhardt,  1946),  western  Montana  (An- 
derson and  Smith,  Hazzard,  1932;  Donaldson,  1934;  Rodgers  and  Jellison, 
1942;  Shpp  and  Carl,  1943),  British  Columbia  (Ricker  and  Logier,  1935;  Slipp 
and  Carl,  1943;  Carl,  1945). 


Map  4 

Habitat:  These  "toads"  live  in  forested  sections,  under  rocks  in  perennial, 
usually  swift-flowing,  small  mountain  streams  of  low  temperature.  After 
heavy  rains  several  collectors  have  found  them  in  moist  woods,  at  varying  dis- 
tances from  the  streams. 

Size:  Adults,  i %-2  inches.  Males,  length  to  tail  29-42  mm.,  tail  3-10  mm. 
Females,  28-51  mm. 

General  appearance:  This  small  "toad"  is  gray,  pink,  or  brown  to  almost 
black.  Numerous  black  spots  occur  on  the  tops  of  the  legs  and  on  the  back. 
The  bar  across  the  head  is  pale  green  to  pale  yellow;  the  underside  of  legs 
rose.  The  females  are  usually  lighter  than  the  males.  The  brown  parotoid 
gland  is  well  developed  or  may  be  broken  into  a  glandular  ridge  along  the 


io6  HANDBOOK  OF  FROGS  AND  TOADS 

side.  The  fingers  are  long  and  slender,  free  of  webbing;  the  toes,  slightly 
webbed.  The  head  is  flattened,  slightly  broader  than  long,  the  snout  obtusely 
pointed,  and  with  no  visible  tympanum.  The  skin  is  smooth  or  slightly  rough- 
ened with  granules,  wrinkles,  warts,  and  small  tubercles.  The  conspicuous 
"tail"  of  the  male  is  level  with  the  ventral  side  of  the  body.  This  is  %-%  inch 
(3-10  mm.)  long,  %-%  inch  (4-5.5  mm.)  broad.  The  anus  is  a  large  swollen 
orifice  just  ahead  of  the  constricted  tip.  The  female  appears  much  like  a  tree 
toad. 

Color:  Carbon  River  Valley,  Rainier  National  Park,  from  J.  R.  Slater,  May 
16,  17,  1930.  Female.  Middle  of  back  from  eye  bar  to  rump  grayish  olive  or 
citrine  drab.  Parotoid  wood  brown,  cinnamon,  or  mikado  brown.  From  paro- 
toid  along  side  is  pecan  brown,  mikado  brown,  orange-cinnamon,  or  sayaFs 
brown.  Same  wash  on  upper  parts  of  tibia  and  an  area  near  anus.  Vertical  part 
of  snout  except  central  bar  (like  bar  between  eyes)  black,  aniline  black,  or  dull 
purplish  black.  Area  below  eye  ageratum  violet  or  vinaceous-purple.  Bar  from 
snout  to  eye  and  behind  eye  to  shoulder  insertion  same  color  as  vertical  snout. 
On  top  of  forelegs  and  hind  legs  and  along  side  a  few  larger  black  spots.  Along 
sides  these  arc  more  elongate  below,  defining  the  browns.  Color  of  hind  legs 
and  forelegs  same  as  mid-back.  Bar  across  head  at  front  of  eyes  glass  green  or 
deep  seafoam  green.  Same  color  on  side  of  face  below  black  nostril,  eye,  and 
shoulder  insertion  bar.  This  bar  bordered  above  with  same  color.  Top  of  snout 
mignonette  green  or  ecru-olive.  Underside  of  body  except  belly  and  femur 
dull  citrine,  ecru-olive,  or  barium  yellow  with  interstices  of  vinaceous,  dark 
anthracene  violet  or  dark  maroon  purple.  Some  of  belly  light  orange-yellow 
with  some  vinaceous  or  pinkish  vinaceous  interspersed.  Femur  pure  deep 
vinaceous  or  purplish  vinaceous.  Ins:  bar  across  top  of  vertical  pupil  cream 
color  or  pale  chalcedony  yellow  with  some  reticulum  of  burnt  sienna  or  hazel. 
This  color  in  upper  part  of  eye  obscures  the  cream  or  yellowish.  Lower  eye 
auburn,  chestnut,  or  chestnut-brown  with  little  specks  of  orange-rufous. 

Male.  Back  kaiser  brown,  hazel,  tawny,  cacao  brown,  or  vinaceous-russet. 
Top  of  hind  legs  and  forelegs  with  numerous  black  spots,  so  also  back;  bar 
from  orbit  to  orbit  pale  chalcedony  yellow.  Top  of  snout  deep  olive-buff.  Ver- 
tical snout  bar  like  eye  bar.  Male  has  venter,  all  except  femur,  like  female  de- 
scribed. Two  patches  on  arm  and  base  of  finger  black  or  seal  brown  or  warm 
blackish  brown.  Iris:  see  female. 

Structure:  The  tail  is  an  intromittent  organ;  lower  jaw  from  below  an  al- 
most perfect  semicircle;  second,  third,  and  fourth  vertebrae  bearing  short  ribs; 
tongue  attached  by  broad  surface,  and  cannot  be  protruded;  arms  of  male  con- 
spicuously heavier  than  in  the  female,  as  also  are  the  hind  limbs  and  feet. 
Breeding  males  with  dark  excrescences  on  the  inner  edges  of  the  inner  fingers 
and  along  the  inner  edge  of  the  forearm,  a  ball-like  excrescence  at  the  base  of 
the  thumb,  and  often  dark  excrescences  on  each  side  of  the  breast;  tuberculcs 


RIBBED  TOADS:  ASCAPHIDAE 


107 


much  finer  on  the  rear  portion  of  the  back  and  on  the  hind  limbs  in  the  fe- 
male; eye  large  with  a  vertical  pupil. 

Voice:  No  vocal  sac. 

Breeding:  Known  dates  are  in  May,  June,  July,  August,  and  September. 
The  eggs  in  circular  masses  of  rosarylike  strings  are  attached  to  the  underside 
of  stones  in  creeks.  The  eggs  are  not  pigmented,  few  in  number,  and  very 
large,  %  inch  (8  mm.),  the  yolk  %  inch  (5  mm.)  in  diameter.  The  mature 
tadpole  is  medium  in  size,  i%-2%c  inches  (42-55  mm.),  its  body  round,  its 
tail  long,  its  crests  not  conspicuous.  The  tooth  ridges  are  %o  (upper  ridges 
with  more  than  one  row  of  teeth).  The  period  of  development  is  probably 
over  winter.  They  transform  during  July,  August,  and  September  at  %-% 
inch  (14-18  mm., possibly  20  mm.).  Wallace  Wood  (Calif.,  1939,  p.  no)  found 
tadpoles  in  January.  Ricker  and  Logicr  (B.C.,  1935,  p.  46)  in  their  five  British 
Columbia  streams  found  one-year  tadpoles  29-41  mm.  and  a  second-year  tad- 
pole 42  mm. 

Journal  notes:  March  16, 1942,  Russ  Grove,  northern  Humboldt  Co.,  Calif. 
Mr.  Milne  says  Klamath  River  is  the  line  of  separation  for  many  forms  such 
as  Ascaphus. 

March  17.  Went  to  south  side  of  Wilson  Creek,  Del  Norte  Co.,  Calif.,  8  miles 
north  of  Klamath  and  close  to  the  ocean,  according  to  directions  of  Wallace 
F.  Wood.  With  seine  and  hoe  we  got  plenty  of  Ascaphus  tadpoles  and  larvae 
of  Rana  boylii  boyht  and  Dicamptodon  ensattis.  There  were  no  adult  frogs  or 
salamanders  under  stones  m  creek  or  under  outlying  ones  or  boards.  When 
you  pick  them  up,  the  Ascaphus  larvae  attach  themselves  to  the  finger  or  net. 
Caught  against  the  side  of  the  can,  the  yellowish  white  tip  with  black  crescent 
band  behind  it  is  conspicuous. 

March  20.  Went  up  MacKenzie  River  watershed  with  Ruth  E.  Hopson  of 
Springfield,  Ore.  .  .  .  At  last  caught  one  Ascaphus  trnei  tadpole. 

March  23.  With  Kenneth  Gordon  up  Santiam  River.  .  .  .  Sixteen  and  one- 
half  miles  from  Foster  at  mouth  of  Moose  Creek.  Storm,  Livczey,  and  I 
pulled  seine  in  creek  for  Ascaphus.  Didn't  get  any,  but  they  are  here.  An  old 
student,  Dr.  A.  C.  Chandler,  secured  them  in  Santiam  National  Forest  25 
years  ago. 

March  29.  Near  Lake  Cushman  dam;  couldn't  get  very  close  because  of  war 
regulations.  Made  a  long  loop  back  to  the  highway  through  the  lumbered 
countryside.  Oh,  the  desolation  of  that  Hamma  Hamma  River  country!  Dr. 
H.  T.  Gaige's  historic  collecting  site  is  under  many  feet  of  water. 

March  31.  With  J.  R.  Slater.  Went  in  afternoon  to  Carbon  River  just  outside 
Rainier  Park  entrance.  In  a  tumbling  hillside  cataract  we  saw  little  A.  truei 
tadpoles  attached  to  rocks;  their  tails  were  wagging  in  the  current.  Caught  no 
adults.  .  .  .  Drove  up  to  ranger  cabin  but  soon  came  back  and  tried  another 
little  stream  which  also  has  Ascaphus  tadpoles. 


io8  HANDBOOK  OF  FROGS  AND  TOADS 

Authorities'  corner:  Mrs.  H.  T.  Gaige  (Wash.,  1920,  pp.  2-3),  working  in 
the  Olympic  region  of  Washington  wrote:  "It  was  under  the  rocks  in  these 
little  creeks  that  Ascaphus  lived.  .  .  .  One  found  them  only  by  working 
slowly  upstream  and  turning  over  every  movable  stone.  Usually  they  either 
floated  down  the  stream  with  the  debris  which  was  released  when  the  stone 
under  which  they  were  resting  was  overturned,  in  which  case  it  took  quick 
action  to  catch  them  before  they  were  out  of  sight,  or  they  made  no  effort  to 
move,  and  in  the  shifting  lights  and  shadows  their  color  so  closely  resembled 
a  bit  of  fir  bark  or  the  small  red  stones  which  were  abundant  in  the  streams 
that  they  were  distinguished  with  difficulty.  Occasionally  they  were  alert  and 
slipped  away  like  a  shadow.  When  placed  on  land  they  were  awkward  and 
stupid  in  action  and  appearance  and  made  little  effort  to  escape.  They  were 
solitary;  never  more  than  one  was  found  under  a  single  stone  and  individuals 
were  usually  well  separated  in  the  stream." 

"The  temperature  of  the  water  at  i  P.M.  registered  10°  Centigrade.  Here 
the  tadpoles  were  very  abundant,  for  as  many  as  seven  were  seen  and  collected 
from  an  area  that  could  be  covered  by  the  palm  of  one's  hand.  They  were  first 
observed  in  the  clear  water  attached  to  the  rocks  on  the  bottom  of  the  creek. 
Efforts  made  to  capture  them  caused  them  to  release  their  holds  and  swim  up- 
stream for  a  few  inches  and  there  seek  shelter  under  the  rocks.  Here  they  were 
more  difficult  to  see  since  their  coloration  blended  nicely  with  the  debris  under 
the  rocks,  but  once  one  knew  where  to  look  and  what  to  look  for,  they  were 
more  obvious"  (A.  and  R.  D.  Svihla,  Wash.,  1933,  p.  38). 

"Trip  July  14-15,  1941.  The  tadpoles  were  at  approximately  the  same  stage 
of  development  as  those  taken  on  June  22,  1930  (which  are  now  in  the  U.S. 
N.  M.).  They  have  short  hind  legs.  The  black  caudal  area  and  milk-white  tail- 
tip  are  prominent  in  most  of  the  individuals. 

"I  watched  one  tadpole  feeding  on  a  stone  over  which  the  swiftest  current 
in  a  riffle  was  falling.  He  moved  about  readily  when  he  desired,  a  millimeter 
or  two  at  a  time,  by  a  slight  'inching  along'  movement  of  the  adhesive  mouth- 
parts.  He  would  turn  this  way  and  that,  his  tail  fluttering  and  flapping  down- 
stream in  the  current,  and  the  slight  but  constant  movement  of  his  mouthparts 
indicated  that  he  was  scraping  at  the  surface  of  the  rock.  Although  nothing 
could  be  seen  on  the  stone,  there  was  probably  a  growth  of  minute  algae.  At 
times  the  tadpole  was  almost  half  out  of  water,  and  again  he  would  browse 
along  till  he  was  nearly  on  the  underside  of  the  stone.  All  of  the  tadpoles  seen 
were  in  the  swiftest  current  available,  usually  on  the  side  of  a  stone  where  a 
drop  of  a  couple  of  inches  formed  a  riffle"  (G.  S.  Myers,  Calif.,  1943,  p.  126). 

"Herewith  are  being  sent  to  you  specimens  of  the  larvae  of  Ascaphus  truei 
that  I  collected  during  the  past  summer.  These  specimens  were  collected  July 
30,  1946  near  Emida,  Idaho.  They  are  from  Township  43  North,  Range  2 
West,  Section  13,  East  Fork  of  Charley  Creek. 

"This  area  is  mostly  in  the  classification  of  open  reproduction,  having  been 


RIBBED  TOADS:  ASCAPHIDAE 


109 


burned  several  times  in  the  past.  The  streams  have  numerous  beaver  dams. 
These  particular  specimens  were  gathered  where  the  water  was  free  flowing 
and  the  stream  bed  of  gravel.  They  were  common  lying  on  the  top  of  the 
gravel  and  offered  no  resistance  to  capture.  I  was  unable  to  locate  any  adults. 
".  .  .  The  boys  in  the  camps  at  first  enjoyed  swimming  in  one  or  another 
of  the  beaver  ponds,  but  they  soon  began  to  complain  of  the  'leeches'  which 
attached  themselves  to  their  legs  and  toes.  I  found  that  these  were  the  same 
larvae.  The  ponds  accumulated  a  silt  deposit  but  were  still  water"  (letter  from 
R.  P.  Ehrhardt,  Kenyon  College,  Ohio,  Sept.  6, 1946). 


no 


HANDBOOK  OF  FROGS  AND  TOADS 


Plate  XVI.  Ascaphus  truei  (XO- i>5-  Fe- 
males. 2.  Tadpole.  3,4.  Males. 


Plate  XVII.  Scaphiopus  couchii.  1,6. 
Males  (X%).  2.  Male  (Xtt).  3-  Female 
(X%).  4-  Male  croaking  (X%).  5- 


FAMILY  SCAPHIOPODIDAE 
Genus  SCAPHIOPUS  Holbrook 

Maps  5-7 

Couch's  Spadefoot,  Southern  Spadefoot,  Rain  Toad,  Sonoran  Spadefoot, 
Sonora  Spadefoot,  Cape  St.  Lucas  Spadefoot 

Scaphioptts  coitchii  Baird.  Plate  XVII;  Map  5. 

Range:  Texas  to  Arizona  and  Utah,  northern  Mexico  and  Lower  California. 

Habitat:  These  spadcfoots  live  in  subterranean  burrows,  often  under  logs  or 
similar  shelter,  and  are  nocturnal  in  habit.  They  breed  in  temporary  pools, 
coming  out  only  after  heavy  rains. 

Size:  Adults,  1%-$%  inches.  Males,  48-70  mm.  Females,  50-80  mm. 

General  appearance:  The  short,  fat,  toadlike  body  has  a  greenish  back,  more 
or  less  marbled.  The  skin  is  roughly  tuberculate,  with  many  light  tubercles  on 
the  sides.  Both  the  ear  and  parotoid  glands  are  indistinct.  The  eyes  are  large 
and  protuberant  with  vertical  pupils.  The  venter  is  whitish;  the  fingers  and 
toes  are  light;  and  the  rear  of  the  arm  and  leg  has  a  light  band.  The  outer  sole 
tubercle  and  sometimes  the  tips  of  fingers  and  toes  are  dark. 

Color:  Male.  Beeville,  Tex.,  March  26, 1925.  Upper  parts  yellowish  oil  green; 
on  sides  same,  oil  yellow  mixed  with  yellowish  oil  green;  tubercles  on  side, 
posterior  back,  and  tibia  calliste  green;  tubercles  below  vent  white;  rear  of  legs 
and  groin  parrot  green;  below  eye  and  lower  muzzle  buffy  citrine;  tympanum 
oil  yellow  or  yellowish  oil  green  and  a  little  livid  brown.  Under  parts  white 
except  small  pectoral  patch  of  Martius  yellow  or  pale  greenish  yellow,  and  area 
from  groin  to  groin  across  rear  of  belly  where  only  livid  brown  or  purplish 
vinaceous  exists.  Eye:  pupil  vertical,  rim  lemon  yellow,  iris  lemon  yellow  to 
bright  green-yellow  reticulated  with  black. 

Female.  Comfort,  Tex.,  May  19,  1925.  Bands  on  front  and  rear  of  upper 
eyelid  citron  green  or  oil  yellow;  same  color  on  side  of  upper  jaw,  above  and 
below  tympanum,  along  sidfcs,  on  back,  on  dorsum  of  hind  limbs,  in  each 
case  interspersed  with  oil  green.  These  two  colors  are  sharply  contrasted  in  the 
female,  while  in  the  male  the  color  is  a  more  uniform  calla  green  or  oil  green, 
with  little  spotting  of  lighter  color.  Black  bars  on  the  limbs,  and  some  black 
on  nostril.  Another  has  mignonette  green  on  sides  spotted  with  black  and  ivy 
green,  tea  green,  or  water  green  on  back.  Fingers  white  on  dorsum,  toes  with 


112 


HANDBOOK  OF  FROGS  AND  TOADS 


some  white  on  clorsum.  Outer  bands  on  feet  whitish.  Under  parts  white  except 
lower  belly  and  under  legs,  which  are  slate  violet  (2)  to  vinaceous-lilac.  Iris 
green-yellow  or  bright  green-yellow  with  black  on  outside,  which  is  sometimes 
a  horizontal  bar  in  front  and  behind.  From  black  on  outside,  black  lines  run 
almost  to  the  vertical  pupil.  In  general  one  can  identify  females  because  they 
are  spotted,  while  males  are  more  uniform. 

Structure:  No  pectoral  gland;  skin  on  crown  of  head  thin;  tongue  subcir- 
cular,  slightly  emarginate;  hand  nearly  as  long  as  forearm;  toes  fully  webbed. 

Voice:  The  chorus  is  harsh  and  noisy,  a  great  caterwauling,  and  can  be  heard 
a  considerable  distance.  The  individual  call — tvow,  me  ow,  or  a  ow — is  a  most 
unearthly  noise  like  that  of  someone  in  pain. 


Map  5 

"The  cry  of  the  male  is  a  loud,  resonant  'ye-ow'  repeated  at  intervals;  that 
of  the  female,  a  short  grunting  'ow1  uttered  several  times  in  shorter  intermis- 
sions11 (J.  K.  Strecker,  Jr.,  Tex.,  igoSe,  p.  203). 

"In  the  case  of  S.  cottchii  it  [the  song)  is  a  bleat  lasting  from  5-6  seconds 
ordinarily.  This  can  be  best  described  as  sounding  so  much  like  the  bleat  of 
a  lamb  as  to  be  mistaken  for  it.  The  song  is  given  while  the  male  sits  upon  the 
mud  near  the  edge  of  the  puddle"  (A.  I.  Ortenburger,  Ariz.,  1925,  p.  19). 

Breeding:  The  time  is  from  April  to  August  at  periods  of  heavy  rainfall. 
The  eggs  are  in  bands  Vi  inch  (6  mm.)  across  or  cylindrical  masses  on  plant 
stems,  the  jelly  is  rather  firm,  the  eggs  are  close  together,  black  above  and 
creamy  white  below.  The  vitellus  is  Ms-He  inch  (1.4-1.6  mm.).  In  warm 
spots  they  hatch  in  iH  days.  The  "bronzy"  tadpole  is  black,  dotted  with  old 


SPADEFOOTS:  SCAPHIOPODIDAE  113 

gold  or  fawn,  small,  i  inch  (24.5  mm.),  broad  with  tail  tip  rounded.  The 
tooth  ridges  are  %,  rarely  %,  %,  %,  %.  After  a  period  of  15-40  days,  the  tad- 
poles transform  during  the  summer  months  and  early  fall  at  %o-%  inch  (7.5- 
12.5  mm.). 

Journal  notes:  May  29,  1925,  Comfort,  Tex.  The  spadefoots  are  calling  as 
they  float  spread  out  on  the  surface.  Their  sides  are  swelled  out  and  vibrating. 
They  often  seem  to  curve  their  backs  in  their  tremendous  efforts. 

June  4.  In  Comfort,  Tex.,  in  roadside  pools  where,  on  May  29,  we  found 
Couch's  spadefoot  breeding,  there  are  now  large  tadpoles.  The  rain  came  May 
28.  The  eggs  must  have  been  laid  then.  How  fast! 

June  7.  At  Encinal,  Tex.,  roadside  pool  alive  with  spadefoot  tads.  .  .  . 
More  spadefoots  2  miles  beyond  Cactus.  Whenever  the  pools  begin  to  dry  up, 
grackles  go  there  to  eat  the  tads. 

June  8.  Near  Dolores,  Tex.,  we  stopped  beside  a  long  roadside  rain  pool.  It 
is  a  sandy  area  with  scattered  bushes,  very  little  herbaceous  material,  and  mes- 
quite  rather  far  apart.  All  stages  were  here,  even  tiny  transformed  ones  leaving 
the  pond.  They  were  hopping  out  so  thickly  that  they  formed  a  seething  mass, 
several  spadefoots  deep.  They  were  gathered  around  small  herbs,  small  bushes, 
and  larger  ones,  when  possible,  for  shade,  and  generally  just  enough  in  a  place 
to  match  the  shadow  of  the  plant.  Many  of  them  crawl  out  of  the  pond  when 
they  still  have  very  long  tails.  I  wonder  if  this  hastens  the  shrinking  of  the  tail. 

June  IT,  east  of  Hebronville,  Tex.  Last  night  we  went  out  with  flashlights. 
Near  a  clump  of  cactus  saw  a  frog.  It  proved  to  be  Couch's  spadefoot.  Caught 
it.  The  instant  we  released  it,  it  started  for  a  clump  of  prickly  pear  and  went 
down  into  a  pack  rat's  hole.  I  verily  believe  this  is  one  place  where  they  aesti- 
vatc. 

Authorities9  corner: 

J.  K.  Strecker,  Jr.,  Tex.,  19086,  p.  203.         F.  W.  King,  An/,.,  1932,  p.  175. 
A.  I.  Ortenburger,  Ariz.,  1925,  p.  20.         L.  W.  Arnold,  Ariz.,  1943,  p.  128. 

Hammond's  Spadefoot,  Western  Spadefoot,  Hammond's  Spea,  Western 
Spadefoot  Toad,  New  Mexican  Spea 

Scaphiopus  hammondn  hammondn  Baird.  Plate  XVIII;  Map  6. 

Range:  California  to  Lower  California,  cast  through  Arizona,  southwest 
Colorado,  New  Mexico,  to  Texas  and  Mexico. 

Habitat:  They  live  underground  in  burrows  which  they  dig  in  soft  earth 
by  backing  into  the  ground  and  digging  with  their  hind  feet,  which  are  armed 
with  spades.  They  rock  the  body  as  they  dig,  and  the  dirt  falls  into  the  bur- 
rows on  top  of  the  toads.  They  breed  commonly  in  temporary  rain  pools  or 
temporary  overflow  areas. 

Size:  Adults,  i%-2%  inches.  Males,  37.5-59  mm.  Females,  37.5-61  mm. 

General  appearance:  The  body  is  stout  and  toadhke,  small  in  size.  The  eyes 


HANDBOOK  OF  FROGS  AND  TOADS 


are  large  and  protuberant  with  vertical  pupils.  The  skin  is  fine,  relatively 
smooth,  dotted  with  fine  roundish  tubercles.  The  back  is  greenish,  the  sides 
yellowish  glaucous,  light  mineral  gray,  or  greenish.  There  are  green  spots  on 
the  back,  top  of  head,  and  legs.  The  forward  under  parts  are  white,  sometimes 


M- 

:      '.V.V    IT    - 


Map  6 

buffy  on  the  throat;  the  rear  under  parts  purplish.  The  males  have  a  wash  of 
grayish  green  on  each  side  of  the  throat. 

Color:  "Color  (in  life)  above  dark  green,  with  scattered  spots  of  dusky;  four 
(or  two)  incomplete  longitudinal  stripes  of  dull  white,  inner  pair  in  line  with 
inner  margins  or  orbits,  the  outer  in  line  with  tops  of  tympanic  membranes; 
tubercles  on  back  and  sides  tipped  with  red  or  orange  in  young  individuals; 


SPADEFOOTS:  SCAPHIOPODIDAE  115 

ventral  surface  plain  white;  throat  region  (vocal  sac)  blackish  in  males,  dusky 
in  females.  Females  lack  the  conspicuous  longitudinal  dorsal  streaking  of 
white,  this  being  broken  and  much  less  extensive"  (T.  I.  Storcr,  Calif.,  1925, 
p.  149). 

Structure:  Head  broader  than  long,  muzzle  short  and  overhanging  the 
lower  jaw;  epidermis  on  top  of  head  thick  and  horny;  tympanum  indistinct; 
no  parotoid  gland;  tongue  very  large,  entire;  no  tibial  or  pectoral  gland;  hind 
limb,  tibia,  foot,  fourth  toe,  and  fingers  relatively  longer  than  in  Couch's  or 
in  Holbrook's  spadefoot. 

Voice:  The  males  call,  lying  on  the  surface  of  the  water.  The  call  is  a  rolling 
or  bubbling  one,  a  croak  more  like  the  croak  of  some  frog  than  Couch's  or  the 
hermit  spadefoot.  It  has  been  described  in  widely  different  ways:  as  the  loud 
purr  of  a  cat  with  the  metallic  sound  of  grinding  gears,  as  a  low-toned  ttrr-r-r-r, 
as  a  loud  crah-crah-t  ah,  and  as  a  resonant  ye-ow.  It  has  been  called  unusual, 
weird,  plaintive,  and  ventriloqual. 

Breeding:  They  breed  from  mid-February  to  August,  dependent  upon 
heavy  rainfall.  The  eggs  are  in  cylindrical  masses  attached  to  grass  or  plant 
stems.  The  eggs  on  the  periphery  of  a  jelly  cylinder  may  look  stalked,  the  stalks 
%-%  inch  (5  or  6  to  9  mm.)  long  and  Vio-/io  inch  (1.4-2.3  mm.)  in  diame- 
ter, the  eggs  Msn-Mo  inch  (1.0-1.6  mm.).  The  eggs  hatch  in  i%-2  days.  The 
dark  greenish  black  tadpoles  may  grow  large,  2% -2%  inches  (65-70  mm.) 
long.  They  are  broad,  almost  round-bodied  in  dorsal  view,  the  eyes  close  to- 
gether, the  tail  short  with  rounded  tip,  the  spiracle  low,  almost  ventral.  Like 
most  spadefoot  tadpoles,  the  musculature  of  the  tail  stands  out  very  promi- 
nently. The  tooth  ridges  are  %,  %,  %,  %>  %•  After  30-40  days  the  tadpoles 
transform  from  May  20  to  September  i,  at  M>-i!4  inch  (13-32  mm.).  The 
tadpole  is  carnivorous  in  habit  and  may  prey  on  its  own  kind,  but  it  is  a  very 
effective  enemy  of  the  mosquito. 

Ortenburger  (Ariz.,  1925,  p.  19)  wrote:  "(a)  Diameter  of  egg  including 
gelatinous  layers — S.  couchit  2.5-3.5  mm.;  S.  hammondn,  1.5-2.0  mm.; 
(b)  thickness  of  jelly  layers — S.  conchii,  usually  more  than  i  mm.;  S.  ham- 
mondit,  mostly  less  than  0.5  mm.;  (c)  method  of  attaching  the  individual  eggs 
to  others  in  mass — S.  cotichh  attached  with  very  short  stalk  and  S.  hammondn 
by  a  slender  stalk  5-10  mm.  in  length  .  .  .  ;  the  eggs  of  S.  hammondti  were 
arranged  on  similar  objects  but  arranged  spirally  around  them,  thus  differing 
from  S.  cottchii  in  which  no  spiral  arrangement  could  be  easily  made  out." 

Our  measurements  closely  follow  those  of  Ortenburger.  The  egg  masses  on 
stems  may  be  from  6  or  7  to  25  mm.  in  diameter.  Egg  stalks  may  be  from  5-9 
mm.  long  and  1.4-2.3  mm.  in  diameter;  vitellus  1.0-1.6  mm.;  hatching  period 
i%-2days. 

Journal  notes:  July  8, 1917.  Quite  a  rain  fell  near  Sierra  Blanca,  Tex.  At  7 
o'clock,  we  heard  no  notes  in  the  creek,  but  later  from  our  camp  one-half  mile 
away,  we  heard  the  chorus  plainly  and  decided  it  must  be  spadefoots.  We 


ii6  HANDBOOK  OF  FROGS  AND  TOADS 

found  toads  and  spadefoots  of  two  species  migrating  from  the  mountain  side 
of  Sierra  Blanca  toward  the  pool  and  noise.  .  .  .  Along  the  edges  of  the  swift 
stream  now  flowing  across  the  flooded  area  we  found  Scaphioptts  couchii. 
Their  cries  were  catlike.  The  5.  hammondii  were  on  the  surface  of  the  water 
and  their  calls  were  bubbling.  .  .  .  The  Hammond's  male  will  float  like 
S.  holbroofyi.  When  he  croaks,  the  rear  half  of  the  back  dips  beneath  the 
water. 

July  9.  The  stream  has  disappeared;  it  is  now  broken  up  by  intermediate 
mud  flats.  The  spadefoots  and  toads  have  disappeared  from  last  night's  ren- 
dezvous. 

July  27,  1925.  Found  Hammond's  spadefoot  eggs,  tads,  and  transformed  in 
ponds  by  the  roadside  16  miles  east  of  Vail,  Ariz.  In  one  pond  must  have  been 
40-50  masses  of  eggs  on  the  ends  of  grass  stems.  The  eggs  seemed  to  be  out 
on  periphery  of  the  jelly  cylinder.  They  look  to  be  stalked.  This  is  the  same 
thing  we  found  in  Texas  Pass  8  years  ago,  when  we  suspected  it  might  be 
Hyla  arenicolor,  but  now  we  know  it  is  S.  hammondii.  These  eggs  were  laid 
last  night  and  are  already  nearing  hatching. 

April  20,  1942.  With  Hadsell  and  Culbertson  to  Fresno,  Calif.,  slough.  Be- 
fore we  reached  White  Bridge  we  came  to  two  or  three  ponds  where  we  took 
large  light-colored  tadpoles  (S.  h.  hammondii).  Here  Culbertson  took  some 
that  were  almost  transformed,  earlier. 

June  13,  Mesa,  Ariz.  Went  out  last  night  toward  Florence  Junction.  Heard 
bullfrogs  and  Hammond's  spadefoot  in  a  pool  at  Desert  Wells. 

July  9,  Lakeside,  Ariz.  Tonight,  early  in  the  evening  heard  no  frogs.  After 
I  went  to  bed  about  10  P.M.  heard  a  few  S.  h.  hammondii  in  one  of  the  ponds 
above  us. 

July  10.  Tonight,  heard  a  few  isolated  S.  h.  hammondii  again. 

Authorities'  corner: 

E.  D.  Cope,  N.  Mex.,  1884,  p.  14, 

T.  I.  Storer,  Calif.,  1925,  pp.  156-157. 

F.  W.  King,  Ariz.,  1932,  p.  175. 

Central  Plains  Spadefoot,  Central  Plains  Spadefoot  Toad,  Cope's  Spea, 
Western  Spadefoot  Toad 

Scaphtopus  hammondii  bombijrons  (Cope).  Plate  XIX;  Map  6. 

Range:  Not  fully  defined.  Stejneger  and  Barbour  apparently  consider  the 
form  from  the  Dakotas  to  Oklahoma,  northwestern  Texas,  New  Mexico,  and 
west  to  Idaho  to  be  S.  h.  bombifrons,  but  V.  M.  Tanner  (1939)  considers  those 
west  of  Utah's  east  line,  north  of  northern  Arizona,  and  east  of  Nevada's  west 
line  to  be  Cope's  old  form  inter montanus.  H.  M.  Smith  (Kansas,  1934)  re- 
marks that  the  range  overlaps  with  S.  h.  hammondii  in  Colorado,  western 


SPADEFOOTS:  SCAPHIOPODIDAE 


117 


Plate    XVIII.    Scaphiopus    hammondii 
hammondii  (X%)-  i>M>6*  Males-  3- 
pole.  5.  Transformed  frog.  7.  Eggs. 


Plate  XIX.  Scap/nopus  hammondii  bom- 
bifrons  (X%)-  1,2,3,5.' Females.  4.  Male. 


n8  HANDBOOK  OP  FROGS  AND  TOADS 

Oklahoma,  and  northern  and  western  New  Mexico.  P.  Anderson  (Mo,,  1945) 
recently  added  Missouri  to  its  range. 

Habitat:  "East  of  Colorado  Springs  in  the  low  rolling  hills  is  one  of  these 
areas.  The  soil  is  a  mixture  of  sand,  gravel,  and  loam  and  generally  quite  dry. 
It  is  here  that  adult  spadefoot  toads  are  found  at  depths  varying  from  a  few 
inches  to  several  feet.  ...  It  usually  chooses  soft  ground  in  which  to  burrow. 
With  its  spade-armed  feet  it  pushes  the  soil  aside,  and  by  a  slow  rocking  move- 
ment sinks  backwards  beneath  the  surface  of  the  ground.  The  heavy  skin  of 
the  head  is  probably  used  to  keep  the  burrow  open  in  front  or  to  pack  the 
earth  of  the  walls  of  the  burrow.  The  descending  toad  leaves  no  trace  on  the 
surface  to  indicate  its  course"  (R.  J.  Gilmore,  Colo.,  1924,  pp.  1-2). 

"Along  the  Powder  River  near  Powderville  in  Montana,  on  June  15,  1916, 
while  lying  upon  my  cot,  I  heard  a  curious  rustling  in  the  dry  leaves  about  our 
tent.  Upon  investigation  with  a  flashlight  many  small  spadefoot  toads  were 
found.  They  were  hopping  about  in  the  dry  leaves  which  were  scattered  about 
on  the  sandy  soil.  When  hunted  with  a  flashlight  they  endeavored  to  burrow 
out  of  sight  and  but  a  few  minutes  were  required  for  them  to  entirely  conceal 
themselves.  These  spadefoots  make  circular  holes  in  the  ground  and  yet  in 
sandy  soil  it  is  very  difficult  to  find  the  place  where  they  have  burrowed  down, 
for  in  most  cases  it  seems  as  if  they  had  pulled  the  hole  in  after  them.  After 
the  breeding  season  is  over  they  take  more  pains  in  constructing  their  burrows, 
as  ...  [the  burrows]  are  well  rounded  and  resemble  somewhat  an  earthen 
jar  with  a  narrow  top.  Around  this  opening  there  is  present  some  sticky  mat- 
ter which  may  aid  in  the  ensnaring  of  insects.  I  have  usually  found  this  toad 
most  plentiful  in  sandy  areas,  especially  along  the  banks  of  streams,  though 
they  occur  on  the  elevated  plains  from  Kansas  to  Montana"  (R.  Kellogg, 
Mont.,  1932,  p.  36). 

Size:  Adults  (roughly)  i%-2%  inches.  Males  38-52  mm.  Females  40-57 
mm. 

General  appearance:  Stout;  small;  vertical  pupil;  interorbital  boss.  Grayish, 
reticulated  or  uniform;  white  below  except  throat  of  male.  Two  series  of  dorsal 
stripes  or  these  absent.  Smoother  than  S.  hammondii.  Internasal  distance  i% 
in  eye  to  nares  in  bombtfrons,  i  in  hammondii. 

Color:  Colorado  Springs,  Colo.,  July,  1928,  from  R.  J.  Gilmore.  Female. 
Sides  yellowish  glaucous  or  light  mineral  gray  or  water  green;  back  in  gen- 
eral, top  of  head,  and  hind  legs  vetiver  green,  tea  green,  or  water  green.  Back 
of  each  eye  on  back  is  a  dull  citrine,  water  green,  or  yellowish  glaucous  spot 
edged  with  Lincoln  green  or  deep  grape  green.  On  hind  legs,  rear  of  back,  and 
top  of  head  are  spots  Lincoln  green  or  dusky  olive-green.  Some  of  these  spots 
have  light  centers  which  are  tubercles  (under  the  lens,  dark  olive-buff  in 
color).  Under  parts  are  white  or  cartridge  buff,  particularly  on  the  throat; 
other  parts  are  congo  pink  to  salmon-buff  or  light  ochraceous  buff.  There  are 
two  tubercles  ahead  of  vent  and  also  beside  it  that  are  white  to  light  buff.  Iris 


SPADEFOOTS:  SCAPHIOPODIDAE  119 

pale  greenish  yellow  or  light  green-yellow  unbroken  around  pupil;  rest  with 
black  lines.  Iris  may  be  orange-pink.  A  spot  or  transverse  bar  like  dorsal  spots 
occurs  on  upper  eyelid. 

Male.  The  males  on  either  side  of  throat  have  wash  of  deep  bluish  gray- 
green,  or  light  terre  verte,  or  grayish  blue-green.  [This  from  memory  of  males 
alive  three  days  ago.]  In  alcohol  the  throats  of  male  are  very  bluish  or  plum- 
beous. 

Structure:  Metatarsal  tubercle  rounded;  parotoid  gland  absent.  "A  Scaphio- 
pus  with  rounded,  not  elongated,  inner  metatarsal  tubercles;  tip  of  fifth  toe 
frequently  blackened  and  corneous;  anterior  interorbital  region  swollen,  con- 
vex; parotoid  glands  indistinct,  tympanum  also  usually;  toes  nearly  fully 
webbed,  fingers  very  slightly;  pupil  vertical"  (H.  M.  Smith,  Kan.,  1934, 
p.  427). 

Voice:  "The  call  of  this  toad  is  quite  weird  and  unusual  and  may  be  likened 
to  the  squawk  of  some  animal  when  severely  injured  or  a  resonant  ye-ow. 
Once  heard  this  distinct  call  is  not  likely  to  be  forgotten"  (R.  Kellogg,  Mont., 
1932,  p.  36).  "After  arriving  at  the  ponds  the  male  spadefoot  indulges  in  very 
vigorous  nuptial  song,  which  continues  without  interruption  until  the  mating 
has  been  completed.  The  efTect  has  been  described  as  'weird  plaintive  cries,' 
'hoarse  and  woeful' "  (R.  J.  Gilmore,  Colo.,  1924,  p.  3). 

Breeding:  These  frogs  breed  from  May  to  August  in  the  rainy  season.  The 
amplexus  is  inguinal.  "The  egg  masses  vary  in  size.  Large  masses  contain  200 
to  250  eggs,  smaller  ones  10  to  50.  The  mass  is  attached  to  submerged  vegeta- 
tion, or  to  any  object  protruding  from  the  bottom.  The  mass  is  elliptical  in 
shape."  "The  incubation  period  as  observed  in  the  field  seems  to  be  less  than 
forty-eight  hours."  "Two  and  one-half  inches  is  the  maximum  length  of  the 
majority  of  adults  in  any  tadpole  community."  "In  1921,  specimens  were 
found  completely  transformed  after  thirty-six  to  forty  days"  (R.  J.  Gilmore, 
Colo.,  1924,  pp.  4,  5).  The  tadpole  is  carnivorous  and  herbivorous. 

From  conversations,  letters,  material,  and  experience,  it  is  evident  that  our 
friends,  Gilmore  and  Kellogg,  know  this  form  best.  Dr.  Gilmore  has  used  this 
species  in  classes  for  a  quarter-century  and  Dr.  Kellogg  has  known  it  equally 
long.  In  recent  years  A.  H.  and  M.  S.  Trowbridge  and  A.  N.  Bragg  of  Okla- 
homa have  added  the  most  to  our  knowledge  of  this  frog. 

In  1934  H.  M.  Smith  characterized  the  tadpole  thus:  "Upper  mandible  with 
a  large  median  beaklike  projection,  lower  mandible  with  a  deep  elevated  me- 
dian notch;  a  black,  corneous  toothlike  projection  from  roof  of  mouth;  buccal 
musculature  conspicuous  and  visible  through  skin  as  are  viscera!"  In  1929  we 
had  described  the  tadpole  of  S.  hammondii  with  no  thought  of  separation  into 
its  three  forms. 

In  1942  A.  N.  Bragg  (Okla.)  characterized  the  tadpole  of  S.  h.  bombifrons 
as  having  "jaws  without  a  beak  and  notch;  jaw  muscles  not  overdeveloped." 
In  1941  in  New  Mexico  he  discovered  that  "Smith's  figure  and  description  are 


120 


HANDBOOK  OF  FROGS  AND  TOADS 


of  S.  hammondii,  and  R.  J.  Gilmore  (1924)  and  Wright's  (1929)  are  of  S. 
bombifrons,  although  labeled  the  reverse  of  this." 

Authorities'  corner: 

J.  A.  Tihen  and  J.  M.  Sprague,  Kans.,  1939,  pp.  501-502. 
G.  A.  Moore  and  C.  C.  Rigney,  Okla.,  1942,  p.  78. 

Great  Basin  Spadetoot  Toad,  Western  Spadefoot  Toad 

Scaphiopus  hammondii  intcrmontanus  (Cope).  Plate  XX;  Map  6. 

Range:  Utah,  Nevada,  northern  Arizona,  western  Colorado,  southwestern 
Wyoming,  Idaho  except  extreme  north,  southwestern  Washington,  eastern 
Oregon — a  Great  Basin  form  (after  V.  M.  Tanner).  British  Columbia. 

Habitat:  Canyon  pools;  desert  springs  and  pools,  intermittent  and  perma- 
nent; irrigation  ditches;  stream  edges;  rain  puddles;  water  pockets;  water  de- 
pressions made  by  cattle. 

Size:  Adults  iM»-2%  inches  (40-63  mm.).  Males,  40-59  mm.  Females,  45- 
63  mm. 

General  appearance:  "The  evolution  of  the  subgenus  Spea  seems  to  be  from 
hammondii  through  bombifrons  to  tntermontanus.  In  these  species  there  is  a 
progressive  development  of  the  osseous  parts  of  the  cranium  with  a  closure 
of  the  fronto-parietal  fontanelle  in  practically  all  specimens  of  intermontanus" 
".  .  .  Fairly  rugose."  "Intermontanus  has  a  greater  internarial  distance  than 
either  bombifrons  or  hammondii"  (V.  M.  Tanner,  Gen.,  1939,  pp.  15,  16). 

"They  appear  to  demonstrate  that  bombifrons  and  hammondii  are  in  the 
same  species,  for  there  is  obvious  intergradation  in  every  character.  Moreover, 
it  seems  likely  that  additional  material  from  this  intervening  area  [Nevada] 


Plate  XX.  Scaphtopus  hammondn  tnterwontanus  (X%)-  From  C.  L.  Patch, 
Okanagan  Landing,  B.C.  Adults. 


SPADEFOOTS:  SCAPHIOPODIDAE  121 

would  demonstrate  the  validity  of  a  third  race,  intermontanus"  "Compared 
with  hammondii  this  form  was  supposed  to  be  distinguished  by  larger  size, 
lighter  colors,  and  the  presence  of  the  superior  pair  of  light  lines"  (J.  M.  Lins- 
dale,  Nev.,  1940,  p.  200) . 

Color:  "Their  color  was  yellowish  olive  above,  spotted  with  darker  olive, 
belly  soiled  white,  chin  darker,  sides  of  front  legs  and  feet  gray,  the  hind  legs 
tinted  on  the  under  surface  with  blood  red.  The  eye  was  large  and  very  bril- 
liant, the  iris  brassy  with  fine  black  reticulations.  The  pupil  was  very  sensitive 
to  light.  There  were  two  broad,  grayish  stripes  on  the  back,  and  one  on  each 
side  of  the  body.  The  glands  were  usually  darker  than  the  surrounding  skin, 
and  in  some  examples  scattered  glands  bore  brick  red  caps,  which  with  darker 
rings  appeared  as  ocelli.  Some  specimens  were  lighter  than  others,  and  an  oc- 
casional one  had  a  strong  infusion  of  pale  reddish  brown"  ( J.  O.  Snyder,  Nev., 
1920,  pp.  83-84). 

"The  colour  in  life  was  grayish  green  above  and  more  or  less  mottled;  a 
longitudinal  whitish  stripe  down  each  side  of  the  back  from  behind  orbits  to 
a  little  behind  knees,  and  one  along  each  side  from  behind  ear;  a  short  median 
whitish  stripe  on  posterior  part  of  back  (over  urostyle) ;  a  darkish  blotch  over 
orbit  passing  backward  and  inward,  and  a  darkish  line  from  eye  to  nostril; 
tubercles  on  back  and  sides  tinged  with  orange;  throat  dusky,  belly  white" 
(E.  B.  S.  Logier,  B.C.,  1932,  p.  319). 

Structure:  "No  interorbital  boss  present.  In  some  specimens  of  intermonta- 
nus  there  is  a  glandular  interorbital  elevation  which  resembles  the  true  boss 
found  in  bombijrons.  This  may  be  removed  and  the  true  nature  of  the  skull 
revealed.  .  .  .  Head  width  wider  20.9-22.5  mm.  Body  rugose  or  with  many 
individual  prominences  or  warts.  Color  mottled  whitish  and  black  above;  ven- 
ter whitish;  in  preservative  the  back  becomes  blackish  with  some  white  areas. 
At  times  the  back  is  streaked  with  whitish  lines.  Venter  white.  No  fronto- 
parietal  fontanelle;  interorbital  space  with  prominent  fronto-parictal  bones 
forming  ridges;  ...  in  some  specimens  the  interorbital  space  is  filled  with  a 
glandular  prominence  resembling  the  bombifrons  species;  head  width  22.5 
mm.,  whole  foot  31.2,  confined  in  the  main  to  the  Great  Basin  area"  (V.  M. 
Tanner,  Gen.,  1939,  pp.  11-12). 

Voice:  "Their  appearance  was  at  once  announced  by  a  loud  chorus  which 
differed  markedly  from  that  of  Hyla  or  Rana,  being  in  a  lower  key,  somewhat 
guttural,  and  a  little  rasping.  It  was  entirely  different  from  that  of  Bufo"  (J.  O. 
Snyder,  Nev.,  1920,  p.  83). 

"I  would  describe  the  call  as  a  soft,  though  very  penetrating  \wa\,  low- 
pitched  with  something  of  the  quality  of  the  vibrating  of  a  heavy  rubber 
band"  (W.  Wood,  Utah,  1935,  p.  101). 

"A  very  secretive,  wholly  nocturnal,  but  really  common  species.  Its  call,  a 
loud  crah-crah-rah,  repeated  at  short  intervals,  was  first  heard  at  Bcllevue  late 
in  April  and  again,  following  a  cold  spell,  on  May  5.  Thereafter  it  increased 


122  HANDBOOK  OF  FROGS  AND  TOADS 

in  volume  nightly  until  June  i,  after  which  it  decreased,  stopping  about  June 
15.  Rain  puddles,  overflows  from  irrigation  ditches,  in  fact  every  pool  of  stand- 
ing water  served  as  breeding  places.  Drying  up  of  the  pools  rarely  permitted 
a  development  beyond  the  egg  or  early  larval  stages"  (G.  P.  Englehardt,  Utah, 
1918,  pp.  77-78). 

Breeding:  Records  indicate  breeding  from  April  (Hardy),  May  (Snyder, 
Carl),  June  (Snyder,  Tanner,  Wood),  July.  Transformations  are  from  late 
May  through  June  and  July  to  August  or  September. 

"This,  the  commonest  Carbon  County  amphibian  was  heard  croaking  on 
April  9, 1937;  eggs  were  collected  April  22.  Numerous;  breeding  near  river  at 
Price,  April  27.  Also  found  breeding  in  puddles  of  rain  water,  in  stagnant 
pools,  in  water,  in  alkali  washes,  in  city  reservoirs  and  swimming  pools,  and  in 
sewage  near  Price  River.  Eggs  deposited  in  the  laboratory  hatched  after  four 
days.  Eggs  collected  in  water  containing  algae  were  green  due  to  inclusion  of 
algae  in  the  outer  jelly  while  eggs  deposited  in  muddy  water  included  soil" 
(R.  Hardy,  Utah,  1938,  p.  99). 

"On  the  evening  of  June  2,  1911, 1  happened  upon  a  small  pond  separated 
from  the  water  of  Pyramid  Lake  by  a  narrow  bar.  The  pond  was  but  a  few  feet 
in  width,  and  perhaps  a  hundred  feet  long.  The  water  was  clear  and  slightly 
alkaline  like  that  of  the  lake.  In  it  were  hundreds  of  spadefoots  depositing 
their  eggs  in  masses  one  layer  deep  on  the  upper  surface  of  small  rocks.  The 
eggs  were  not  piled  up  after  the  manner  of  frogs,  nor  were  they  in  strings  like 
those  of  toads.  One  mass  presented  fresh  eggs  and  likewise  others  in  which 
development  was  marked,  plainly  indicating  that  the  mass  was  made  of  at 
least  two  contributions"  (J.  O.  Snyder,  Nev.,  1920,  p.  84). 

Tanner  records  and  describes  a  5i-mm.  tadpole,  coppery  color  in  water, 
bluish  black  in  preservative.  He  gives  labial  teeth  rows  as  %.  Snyder  secured 
a  6o-mm.  tadpole  with  teeth  %,  %  and  transformation  sizes  of  20-24%  mm. 

"From  accounts  of  spadefoots  that  I  have  read  I  judged  that  the  toads  come 
out  of  their  burrows  only  after  a  rain,  and  that  a  certain  amount  of  moisture 
was  necessary  to  bring  them  to  the  surface.  However,  the  fact  that  so  many 
hundreds  of  individuals  came  out  of  the  ground  in  Smoky  Valley  in  1932, 
when  no  rain  had  fallen  for  a  long  time,  indicates  that  other  circumstances  arc 
able  to  initiate  activity  in  this  species."  "In  1933,  with  Compton  I  worked  in 
this  same  vicinity  from  mid-May  to  mid-June.  The  pasture  was  fairly  dry  this 
time,  but  a  large  proportion  of  the  1932  brood  of  spadefoots  had  survived,  and 
we  saw  them  almost  daily.  We  were  surprised  to  see  them  day  after  day  forag- 
ing on  the  surface  of  the  ground  in  daylight  hours"  (J.  M.  Lmsdale,  Nev., 
1938,  pp.  21, 23). 

"On  July  4,  many  small,  recently  transformed  specimens  were  collected  in 
mud  cracks  in  the  bed  of  a  dricd-up  irrigation  ditch  near  Maggie  Creek.  It 
was  found  that  the  easiest  method  of  collecting  them  was  to  stamp  on  the 
ground.  The  vibration  disturbed  them  and  they  would  thrust  their  heads  out 


SPADEFOOTS:  SCAPHIOPODIDAE 


123 


of  the  cracks.  If  the  jar  continued,  they  came  out  and  hopped  about  on  the 
ground  where  they  were  conspicuous  and  easily  captured,  but  when  the 
stamping  ceased  they  soon  disappeared.  A  few  had  completely  transformed, 
but  the  majority  had  tails  varying  in  size  from  mere  rudiments  to  the  length 
of  the  head  and  body.  Later  many  half  grown  specimens  were  observed  com- 
ing up  out  of  the  ground  behind  the  mowers  in  a  hay  field  near  Annie  Creek" 
(A.  G.  Ruthven  and  H.  T.  Gaige,  Nev.,  1915,  p.  16). 

Journal  notes:  May  20, 1942,  Beaver  Dam  Lodge,  Ariz.,  at  bridge.  We  heard 
in  several  directions  a  few  Scaphtopus  hammondii  subspecies.  They  were 
mainly  concentrated  in  the  big  pool  at  north  end  of  bridge.  Two  were  too  far 
out  to  collect — only  one  did  we  get.  Its  call  is  a  harsh  far-carrying  call  very  dif- 
ferent from  the  trill  of  B.  woodhousn. 

June  5.  Stopped  at  Jacob's  Lake,  Ariz.  The  assistant  forester,  Harlen  John- 
son, said  about  May  i  they  heard  an  awful  squawking  and  he  wondered  what 
it  was.  I  asked  him  if  they  had  spadefoots.  He  said  "Yes."  He  sent  droppings 
of  coyotes  to  the  Colorado  laboratory  and  they  said  there  were  spadefoot  re- 
mains in  them.  I  went  down  to  the  lake  and  found  two  sizes  of  Hammond's 
spaclcfoot  tadpoles. 

Authorities9  corner: 
E.  B.  S.  Logier,  B.C.,  1932,  p.  319.    W.  F.  Wood,  Utah,  1935,  pp.  101-102. 

Spadefoot,  Holbrook's  Spadefoot,  Hermit  Spadefoot  Toad,  Spadefoot  Toad, 
Hermit  Toad,  Solitary  Toad,  Solitary  Spadefoot,  Hermit  Spadefoot, 

Storm  Toad 

Scaphioptts  holbroofyi  holbrookji  (Harlan).  Plate  XXI;  Map  7. 

Range:  From  West  Virginia  (Green  and  Richmond,  1942)  and  Ohio,  down 
the  Ohio  River  to  southeastern  Missouri  and  northeastern  Arkansas,  across 
extreme  southeastern  Texas,  and  along  the  Gulf  and  Atlantic  coasts  to  Massa- 
chusetts. 

Habitat:  Shallow  burrows  in  the  ground;  nocturnal  in  habit.  They  breed 
only  after  heavy  rams,  and  then  usually  in  temporary  rain  pools.  The  follow- 
ing note  relates  several  items  about  their  general  habits.  Dave  Mi/zell  of  the 
Folkston,  Ga.,  region,  told  of  "a  frog  called  'storm  frog'  because  it  appears 
when  there  are  storms  or  floods.  It  hollers  wanl(!  It  folds  up  so  that  you  can't 
see  its  legs.  It  is  larger  than  the  'hop  toad.'  One  was  dug  up  in  a  potato  field." 
"Very  common  at  breeding  time;  seen  on  warm  nights  at  the  mouth  of  bur- 
rows, and  hopping  about  in  the  woods"  (O.  C.  Van  Hyning,  Fla.,  1933,  p.  3). 

No  other  form  in  this  country  has  had  more  printed  about  it  than  this  elu- 
sive form.  Our  account,  however,  will  be  one  of  our  shortest  summaries.  No 
one  has  nightly  visited  these  creatures  in  their  nonbreeding  holes  more  than 
Neil  Richmond. 

Size:  Adults,  2-2%  inches.  Males,  54-72  mm.  Females,  50-71  mm. 


I24 


HANDBOOK  OF  FROGS  AND  TOADS 


General  appearance:  Like  the  common  toad,  it  is  short  and  broad  in  body, 
and  with  small  round  posttympanal  glands  (parotoids) .  The  skin  is  relatively 
smooth  but  bears  scattered  warts;  is  usually  brown  in  color,  frequently  with 
two  more  or  less  evident  light  dorsal  stripes.  The  arms  and  legs  are  short  and 
thick;  the  feet,  broad.  The  inner  tubercle  of  the  sole  is  a  large  dark  horny 
process  with  which  the  toad  digs  its  burrows.  The  eyes  are  large  and  protu- 
berant with  vertical  pupils  indicating  nocturnal  habits.  The  throat  and  breast 
are  white;  the  lower  belly  is  grayish. 


Map  7 

Color:  Milliard,  Fla.,  Aug.  18,  1922.  Male.  Stripe  from  eye  back  to  vent 
lemon  yellow  or  greenish  yellow;  other  light  colors  of  the  back  are  same  as 
stripe  from  eye  to  vent,  so  also  on  the  sides.  The  snout  is  mummy  brown, 
Prout's  brown,  sometimes  blackish  brown,  or  bone  brown.  This  is  also  the 
remaining  color  of  the  back,  which  is  sometimes  virtually  black.  Limbs  on 
dorsum,  much  the  same  color  or  slightly  washed  out  greenish  yellow  of  the 
light  back  colors;  under  parts  of  hind  limbs  and  lower  belly  light  grayish 
vinaceous.  Rest  of  the  under  parts  are  white,  especially  the  white  glistening 
throat,  which  shows  so  conspicuously  in  the  "croaking  bubble."  First  three 


SPADEFOOTS:  SCAPHIOPODIDAE 


125 


fingers  with  black  excrescences. 
Spade  of  hind  foot  dark-edged,  as  is 
also  the  tip  of  first  toe.  Web  of  foot 
much  darker  than  the  grayish  vi- 
naceous  web  of  female.  Iris  light 
greenish  yellow  with  black  on  the 
outer  rim.  Pupil  vertical. 

Female.  Sometimes  uniform 
warm  sepia  or  bone  brown  above. 
Throat  and  breast  white.  Underside 
of  fore  limbs  and  hind  limbs  and 
lower  belly  light  grayish  vinaceous. 
Sometimes  the  stripe  on  females 
may  be  sulphur  yellow  instead  of 
the  intense  greenish  yellow  of  the 
male.  Almost  always  each  male  of 
a  pair  had  a  predominance  of  yel- 
lowish hues  and  the  females  in- 
clined to  the  brownish  hues. 

Structure:  Tympanum  much 
smaller  than  the  eye;  large,  wide 
hind  feet;  skin  on  crown  of  head, 
thin;  parotoid  gland  present;  male 
with  a  subgular  vocal  sac;  males 
with  fingers  broader  than  females. 

Voice:  Hoarse,  coarse,  monosyl- 
lable, waftl{,  wanl{,  like  the  calling 
of  young  crows.  Aug.  16,  1922,  we 
heard  the  congress  at  a  half-mile  dis- 
tance. At  this  distance,  to  one  it 
sounded  like  the  calling  or  snarling 
or  complaining  note  of  a  cross  baby; 
to  another  member  of  the  party,  like 
young  crows;  or  at  other  times  like 
young  herons  in  a  herony.  One 
member  characterized  it  as  naarh 
naarh,  complaining,  nasal,  not  shrill 
or  high-pitched.  Near  by  it  sounds 
somewhat  like  where,  where,  where, 
where. 

When  we  first  approached,  the 

males  called  on  all  sides  in  full  daylight,  almost  at  our  very  feet.  The  inflated 
throat  by  day  is  a  beautiful  glistening  white  golf  ball.  The  male  before  he  calls 


Plate  XXL  Scaphtopus  holbroo^ti  hoi- 
brookii.  1,4.  Males  (X%)-  2,3.  Females 
(X%)«  5-  F-gg  mass  (X1/^-  6-  Forearm  of 
male  (X%). 


126  HANDBOOK  OF  FROGS  AND  TOADS 

lies  on  the  water's  surface  with  hind  legs  partially  submerged.  When  he  croaks 
he  dips  the  hind  end  of  the  body  and  the  head  is  reared  to  a  45°-75°  angle 
with  the  water's  surface.  When  at  the  height  of  the  performance,  or  slightly 
before,  he  closes  his  eyes.  Then  the  throat  deflates  and  the  body  inflates.  He 
croaks  about  once  in  every  two  seconds. 

A  big  congress  may  make  as  much  noise  as  a  steam  calliope,  though  of  a 
different  nature.  Normally  the  mated  pairs  are  quiet.  It  is  the  calls  of  free 
males  which  are  searching  a  mate  or  annoying  the  already  mated  pairs  which 
make  up  the  chorus. 

"The  peculiar,  harsh  croaking  of  this  singular  toad  must  be  heard  to  be 
appreciated,  and  can  then  never  be  confounded  with  that  of  any  other  species. 
The  only  sound  we  can  liken  it  to  is  that  of  a  heavily  loaded,  creaking  wagon 
rolling  over  hard  and  uneven  ground"  (F.  W.  Putnam,  Mass.,  1867,  p. 
109). 

"We  have  found  only  one  reference  to  a  spadefoot  singing  while  under- 
ground (A.  H.  Wright,  1932).  We  found  this  to  be  one  of  the  most  striking 
peculiarities  of  the  animal.  In  the  early  part  of  the  evening,  between  nine  and 
ten  o'clock,  no  toads  were  visible  in  the  ponds,  but  from  all  around  the  margin 
came  their  much  muffled  calls.  It  was  a  most  peculiar  sensation  to  be  in  the 
midst  of  the  chorus  but  to  have  only  a  barren  expanse  of  sticky  clay  visible  un- 
der the  flashlight.  Excavating  with  considerable  difficulty  we  found  the  song- 
sters to  be  several  inches  below  the  surface  in  apparently  quite  solidly  packed 
earth.  No  sign  of  entrance  to  the  burrows  was  visible.  As  the  evening  wore 
on  one  after  another  pushed  out  to  the  surface  and  entered  the  pond  to  float 
and  sing  with  greater  vigor.  The  chorus  reached  its  height  shortly  after  mid- 
night" (E.  G.  Driver,  Mass.,  1936,  pp.  67-68). 

Breeding:  On  this  topic  as  on  voice  there  are  countless  observations.  The 
spadefoot  breeds  from  March  to  September  at  periods  of  heavy  rainfall.  The 
eggs  are  in  irregular  bands  along  grass  blades  or  plant  stems,  the  band  1-2 
inches  (25-50  mm.)  wide  and  1-12  inches  (25-300  mm.)  long,  the  egg  1/ic- 
Ki>  inch  (1.4-2  mm.),  the  envelope  %o-/u  inch  (4-5.6  mm.).  They  hatch  in 
i1/i>-2  days.  The  bronzy  tadpole  is  small,  i%  inches  (28  m.)  broad,  but  not 
deep,  its  tail  short  and  rounded.  After  14-60  days,  the  tadpoles  transform  from 
July  to  September  at  %-%  inch  (8.5-12  mm.). 

Journal  notes:  Aug.  16,  1922.  We  started  from  Callahan  in  a  hard  rain,  a 
little  before  noon.  On  a  detour  two  miles  south  of  Hilliard,  Fla.,  we  stopped 
for  cars  going  across  a  swollen  creek.  Francis  went  to  look  for  birds  and  heard 
spadefoots  calling.  We  drove  through  the  woods  and  back  on  the  Dixie  High- 
way and  pitched  camp,  i  mile  south  of  Hilliard  on  an  oak  ridge.  An  old  road 
filled  with  water  made  a  shallow  pond,  and  here  we  saw  the  males  croaking, 
their  white  throats  looking  like  shiny  white  golf  balls.  Just  beyond  was  a 
shallow  surface  pool  made  by  the  heavy  rains.  The  ground  was  covered  with 
herbs:  a  little  Xyris,  a  few  sedges,  Rhexia,  a  small  umbelliferous  plant  with 


SPADEFOOTS:  SCAPHIOPODIDAE 


127 


violet-shaped  leaf,  wire  grass,  and  Hypericum.  The  spadefoots  were  calling 
here,  in  another  similar  pool,  and  in  a  third  deeper  pool  as  well.  At  a  distance 
the  chorus  sounded  like  young  crows  trying  to  call.  The  pond  was  filled  with 
mated  pairs. 

A  few  eggs  had  been  laid.  The  eggs  were  laid  in  more  or  less  irregular  band 
form  along  the  grass  blades  or  plant  stems.  In  the  third  pond  where  the  water 
was  deep,  the  bands  were  long.  The  pair  might  be  floating  on  the  surface. 
When  ready  to  lay,  they  went  to  the  bottom  of  the  pond,  often  the  male  with 
his  eyes  closed  and  the  female  with  hers  partly  closed.  They  moved  along 
slowly  on  the  bottom  or  rested  a  minute.  When  she  found  a  stem  to  suit  her, 
she  seized  it  with  her  front  feet  and  pushed  with  her  hind  feet.  The  male  clung 
close  to  her  back,  his  chin  tight  against  her  back.  (One  we  photographed  had 
an  abrased  chin  as  if  from  pressure.)  He  held  his  knees  against  her  knees,  or 
sometimes  his  feet,  which  are  conspicuously  broad,  were  pressed  against  her 
feet.  She  walked  or  climbed  up  the  grass  blade  or  along  it,  if  it  fell  to  hori- 
zontal position,  and  pressed  her  vent  against  the  blade  as  she  laid  the  eggs.  He 
humped  his  back  to  press  his  vent  close  to  hers  while  she  was  laying.  As  they 
reached  the  top  of  the  blade,  they  sometimes  moved  immediately  to  a  nearby 
one,  or  rested  a  short  period.  When  first  laid,  the  eggs  had  an  irregular  band 
appearance  as  they  were  strung  along  the  blade,  sometimes  being  much 
thicker  if  more  eggs  had  been  emitted  at  such  periods.  When  first  laid,  they 
had  a  brownish  appearance  with  conspicuous  creamy-white  vegetative  pole. 
As  the  jelly  swelled  and  the  eggs  all  turned  right  side  up,  they  looked  very 
black. 

By  the  next  morning  some  eggs  were  almost  ready  to  hatch  (these  must 
have  been  the  ones  we  found  when  we  first  found  the  pond).  The  other  clus- 
ters were  swollen  into  loose,  irregular,  elongate  bunches  attached  to  the  stems 
which  tipped  so  that  many  times  the  bunches  lay  lengthwise  on  the  water. 
There  seemed  to  be  a  tendency  for  the  bunches  of  eggs  to  be  more  or  less  clus- 
tered in  areas.  We  noticed  many  pairs  close  together  that  first  afternoon,  and 
unattached  males  trying  to  get  to  a  female,  thus  making  tangled  masses  of 
toads.  There  was  a  strong  chorus  that  night  and  by  the  next  morning  the  pond 
was  all  churned  up  and  muddy.  Many,  many  eggs  were  there,  but  no  toads. 
The  story  was  told  for  the  season. 

Authorities'  corner: 

A,  Nichols,  Mass.,  1852,  pp.  113-115.          F.  Overton,  N.Y.,  1915,  p.  17. 
F.  W.  Putnam,  Mass.,  1865,  p.  229.  N.  D.  Richmond,  Va.,  1947,  pp.  53-67. 

Key  West  Spadefoot 

Scaphiopus  holbroo1(ii  albits  (Carman).  Plate  XXII;  Map  7. 

Range:  Florida  Keys  and  possibly  the  extreme  southern  part  of  Florida. 
Habitat:  The  query  that  naturally  arises  is:  Do  the  other  species  of  frogs 


128 


HANDBOOK  OF  FROGS  AND  TOADS 


such  as  the  green  tree  frog,  southern  tree  toads,  toads,  or  frogs  have  a  similar 
tendency  toward  albinism  in  the  Keys?  Is  it  due  to  salt  or  a  light  beach  habi- 
tat? Or,  was  this  lot  of  spadefoots  an  isolated  albinistic  collection  that  might 
possibly  occur  on  the  mainland  in  other  portions  of  the  spadefoot's  range? 
One  would  expect  the  subterranean  spadefoot  to  tend  more  toward  albinism 
than  almost  any  other  species  of  the  country. 


Plate  XXII.  Scaphiopus  holbrool{ii  albus  (XO-  i>2>3-  Males,  trom  UbJNM. 
4.  Male,  from  Gainesville,  Fla.  5.  Female,  from  Gainesville,  Fla. 

Size:  Adults  2%-2%  inches.  The  three  specimens  in  the  National  Museum 
are  males  measuring  54,  56,  56  mm.  The  University  of  Michigan  Museum  has 
a  female  56  mm. 

General  appearance:  This  spadefoot  is  like  Holbrook's  but  with  an  exces- 
sive amount  of  white  in  the  pattern. 


SPADEFOOTS:  SCAPHIOPODIDAE 


129 


Color:  "Average  size  less  than  that  of  preceding  (S.  h.  holbroo\ii).  Brown 
of  the  back  lacks  the  red  or  chocolate  tinge.  Readily  distinguished  by  the  great 
amount  of  white  on  back,  flanks,  and  upper  surface  of  limbs.  The  white  forms 
spots  or  vermiculations  which  coalesce  into  bands  of  irregular  shape  and  ex- 
tent" (S.  Carman,  Gen.  Check.  L.,  1884,  p.  45).  A  doubtful  subspecies. 

Structure:  Apparently  the  interorbital  distance  is  narrower  in  S.  h.  albus 
than  in  S.  h.  holbroofyi,  being  in  body  length  9.3-10  in  S.  h.  albus  and  6.77-8.5 
in  S.  h.  holbroo\ii.  Two  pectoral  glands  are  present. 

Scaphiopus  h.                                        albus  holbroo\ii 

Male         Female  Male         Female 

Body  length                               56mm.        56mm.  56  mm.        56mm. 

Interorbital  space                          6.0  mm.       5.5mm.  8.0  mm.       8.0  mm. 

Voice:  Their  cries  sound  like  "ow,  ow,"  and  "miow,"  but  the  latter  much 
deeper  in  tone  than  the  well-known  cat  cry.  The  noise  made  by  a  dozen  males 
is  deafening  when  one  is  near,  though  the  call  lacks  carrying  power  (R.  F. 
Deckert,  Fla.,  1921,  p.  22). 

"The  habits  of  the  two  seem  to  me  to  be  identical,  and  I  can  detect  no  dif- 
ferences in  the  voices"  (A.  F.  Carr,  Jr.,  Fla.,  1940^  p.  54). 

Breeding:  We  have  seen  live  specimens  from  Gainesville  which  seemed 
almost  as  light  as  the  preserved  specimens  of  this  subspecies.  Pending  the  de- 
termination of  the  status  otS.h.  albus  and  whether  it  extends  to  the  southern 
tip  of  Florida,  we  wish  to  point  out  that  Richard  F.  Deckert  does  not  describe 
the  spadefoots  of  Miami  as  S.  h.  albus.  He  calls  them  Scaphiopus  holbrootyi. 

"Scaphiopus  holbroofyi  (Harlan).  On  April  23  (1920)  a  male  specimen  was 
found  by  the  writer,  about  six  inches  down,  in  sandy  marl  at  Brickell  Ave.  and 
Broadway,  Miami.  During  a  prolonged  thunderstorm  many  of  the  spadefoot 
toads  were  encountered  by  the  writer  on  the  streets  south  of  Miami  River,  on 
the  afternoon  of  May  16,  and  during  the  night  were  found  breeding  at  igth 
Street  and  Ave.  H.,  also  at  22nd  Street  and  Miami  Ave.,  and  great  numbers 
were  reported  from  the  low  grounds  near  the  'Alligator  farm/  Miami"  (R.  F. 
Deckert,  Fla.,  1921,  p.  22). 

Journal  notes:  March  18, 1934.  Looking  for  Florida  spadefoots,  we  spent  one 
night  at  Caribee  Colony  on  Matecumbe  Key,  and  went  out  in  the  evening 
with  our  flashlights  to  "shine  their  eyes."  A  few  were  out,  but  more  were  in 
their  burrows  with  the  tops  now  shaken  open. 

Authorities'  corner: 
C.  E.  Burt,  Gen.,  1938,  p.  335. 
V.  M.  Tanner,  Gen,,  1939,  p.  3. 
A.  F.  Carr,  Jr.,  Fla.,  i94ob,  pp.  31,  53-54. 


130 


HANDBOOK  OF  FROGS  AND  TOADS 


Hurter's  Spadefoot,  Hurter's  Solitary  Spadefoot 

Scaphiopus  holbrootyi  hurtcrii  (Strecker).  Plate  XXIII;  Map  7. 

Range:  Eastern  half  of  Texas.  Records  exist  from  Houston  and  Edna  to 
Cameron  County.  Smith  extends  it  to  western  Arkansas. 

Habitat:  Like  other  spadefoots,  these  frogs  come  out  of  their  burrows  to 
breed  in  temporary  pools. 

Size:  Medium.  Type  67  mm.,  from  Waco.  Refugio  specimen  63  mm.  The 
range  of  size  of  ten  breeding  adults  from  Lytle,  Texas  (collected  by  A.  J.  Kirn, 
June  28,  1931)  is  66-78  mm.  Adults,  r%-3%  inches.  Males,  43-73  mm.  Fe- 
males, 44-82  mm. 


Plate  XXIIL  Scaphtopus  holbwolyi  hurtem  (X%)-  i>2»3>4>6.  From  Albert 
Kirn,  Somerset,  Tex.  5.  From  USNM  to  show  pectoral  glands. 

General  appearance:  "Head  short,  length  about  equal  to  width.  (In  hoi- 
brooch  the  head  at  angle  of  mouth  is  much  wider  than  long.)  Snout  heavy 
and  blunt,  not  extending  beyond  the  mouth.  Parotoids  nearly  round,  higher 
and  even  more  conspicuous  than  in  the  eastern  species.  Tympanum  distinct 
but  rather  smaller  than  in  holbroofyi.  (In  type  hardly  more  than  half  the 
diameter  of  the  parotoid.)  Crown  distinctly  rugose.  No  black  granules  in 
space  between  and  in  front  of  the  eyes.  Upper  surfaces  with  small,  closely  set 
tubercles,  very  uniform  in  size  and  distribution.  Many  tubercles  on  sides,  but- 
tocks, and  posterior  portion  of  the  abdomen. 

"Color  above,  pale  greenish,  with  a  pale  yellowish  line  from  each  orbit; 
these  converge  again  in  the  coccyx.  Upper  surface  of  head  and  area  between 
the  light  lines,  dark  plumbeous,  parotoids  olive.  Sides  of  head  and  under  sur- 
faces yellowish-white"  (J.  K.  Strecker,  Jr.,  Tex.,  1910,  pp.  116-117). 

Color:  Somerset,  Tex.,  from  Al  Kirn,  June  20, 1945.  Males.  The  dorsal  back- 


SPADEFOOTS:  SCAPHIOPODIDAE  131 

ground  color  is  Saccardo's  olive,  medal  bronze  to  raw  umber.  A  light  stripe  on 
either  side  crosses  the  rear  of  the  eyelid,  outlines  the  corner  of  the  head  boss, 
the  pair  becoming  parallel  for  %  inch  and  about  %  inch  apart,  next  bowing 
outward  and  then  approaching  each  other  toward  the  groin.  These  stripes  arc 
dull  green-yellow  or  bright  chalcedony  yellow  to  buffy  citrine.  From  the  vent 
forward  is  a  short  middorsal  stripe  of  the  same,  which  also  occurs  on  outer  edge 
of  foot  and  tarsus  and  on  a  few  tubercles  on  the  outer  edge  of  tibia  and  along 
the  sides  of  the  body.  There  is  a  patch  of  the  same  color  above  the  arm  inser- 
tions. The  tympanic  area  is  buffy  citrine  to  dull  green-yellow.  The  top  of  head 
and  boss  are  smooth  and  unspotted  mummy  brown.  The  iris  is  light  cendre 
green  to  light  ochraceous-buff  with  a  horizontal  bar  of  black,  which  is  seldom 
complete.  The  under  parts  are  cream  to  white,  most  intense  on  chin  and  pec- 
toral region.  The  rear  portions  are  vinaceous-buff,  which  color  sometimes  suf- 
fuses the  whole  under  parts.  The  upper  %  of  first  finger  and  inner  and  upper 
half  of  second  bear  black  excrescences. 

Females.  Females  have  a  tendency  to  be  less  green.  The  stripes  are  more 
buffy,  in  one  almost  white.  The  lateral  tubercles  may  be  almost  white  and 
arranged  in  5-7  series.  The  excrescences  on  fingers  arc  lacking. 

Structure:  The  boss  on  head  is  large,  conspicuous,  and  much  raised  at  the 
rear.  The  axillary  breast  glands  are  present. 

"Many  pustules  on  upper  surface  of  tibia.  Glands  on  thorax  present,  con- 
spicuous. Enlargements  resembling  glands  on  inferior  surface  of  femur  (pres- 
ent in  both  specimens).  Spade-like  process  of  foot  narrowly  margined  with 
black.  Palmar  tubercles  rather  small.  Fingers  slender.  Tibia  about  equal  to 
that  of  S.  holbroo^ii  but  femur  and  foot  much  shorter"  (J.  K.  Strecker,  Jr., 
Tex.,  19103,  pp.  116-117). 

Distinguished  from  S.  h.  holbroofyi  by  "its  more  compact  form,  narrow 
head,  blunt  muzzle,  unusually  high  parotoids,  smaller  palmar  tubercles,  and 
shorter  hind  limbs.  The  sides,  buttocks,  tibia,  and  posterior  portion  of  the 
abdomen  are  covered  with  tubercles  instead  of  being  almost  perfectly  smooth. 
The  tubercles  on  the  upper  surfaces  are  more  uniform  in  size"  (same,  p.  116). 

Mr.  Kirn's  material  (June  28-29, 1931),  when  compared  with  male  S.  h.  hol- 
brootyt  of  the  same  size,  has  smaller  measurements.  The  head  to  angle  of 
mouth,  the  width  of  head,  the  tympanum  (equal  in  one),  the  snout  (greater 
in  one)  were  less  than  in  S.  h.  holbrootyi.  The  hind  limbs  were  equal  in  the 
two  species. 

Mr.  Kirn's  specimens  include  66-,  68-,  68-mm.  males.  We  studied  also  Strcck- 
er's  material  and  that  of  USNM  and  Baylor  University.  Our  custom  of  meas- 
uring males  and  females  of  every  species  of  frog  in  the  United  States  at  20, 28, 
36, 44,  56, 66,  68,  and  82  mm.  makes  the  measurements  more  readily  compara- 
ble than  random  ones  of  unscxed  specimens.  The  following  table  is  part  of 
the  25  measurements  we  made  for  each  specimen: 


132  HANDBOOK  OF  FROGS  AND  TOADS 


5.  h. 

S.  h. 

S.  h. 

S.  h. 

hurteni 

holbroo^ii 

hurterii 

holbrool(ii 

44  mm. 

44  mm. 

56  mm. 

56  mm. 

Head  to  angle  of  mouth 

H 

*5 

15 

18 

Width  of  head 

*7 

19 

21 

23 

Snout 

8 

8.5 

9 

ro 

Tympanum 

3 

4.0 

3-5 

5-5 

Hind  limb 

43 

45 

53 

49 

Tibia 

14 

'5-5 

18 

17-5 

Foot  with  tarsus 

22 

24 

28 

28 

Foot  without  tarsus 

16 

16.5 

20.5 

i9 

Voice:  "The  call  is  a  single  note,  which,  while  guttural,  has  a  peculiar  soft 
quality  not  unpleasing  to  the  human  ear,  quite  different  from  that  of  Scaplno- 
pus  bombifrons,  S.  cotichii,  or  5.  hamrnondii,  and  different  also  from  descrip- 
tions of  the  cry  of  S.  h.  holbroolyi,  which  is  commonly  stated  to  be  very  harsh 
and  loud  (cf.  Ball,  1936).  Nevertheless  the  voice  of  hurteni  has  a  quality  char- 
acteristic of  spadefoot  breeding  calls;  I  recognized  it  at  once  when  first  heard 
as  a  spadefoot  call.  The  cry  is  not  exceptionally  loud,  although  a  large  chorus 
can  be  heard  for  at  least  one-half  mile.  (I  have  heard  S.  bombtfrons  for  more 
than  2  miles  on  a  still  prairie  night.)  The  calls  of  males  tend  to  stimulate  other 
males  to  call  and  to  attract  both  females  and  other  males  to  a  given  pool. 

"Each  call  is  explosively  given,  and  the  vocal  sac  becomes  fully  distended 
and  then  deflated  each  time.  Intervals  between  calls  vary,  both  with  the  num- 
bers of  individuals  present  and  with  numbers  calling  at  any  one  time.  The 
usual  interval  m  a  large,  excited  congress  is  from  one  to  two  and  one-half  sec- 
onds. There  is  no  tendency  for  the  calls  of  males  to  be  synchronous"  (A.  N. 
Bragg,  Okla.,  1944,  pp.  231-232). 

Breeding:  January  to  December  (Kirn). 

"All  masses  of  eggs  of  S.  httrtern  seen  were  produced  near  the  shore,  at  and 
near  the  water  surface,  and  strung  out  over  vegetation.  When  in  place,  they 
superficially  resemble  those  of  Bttfo  rather  than  those  of  other  spadcfoots.  Each 
egg  is  black  at  the  animal  pole,  shading  to  very  light  grey  or  white  at  the  vege- 
tal. Each  is  inclosed  in  a  ball  of  jelly  considerably  larger  than  itself,  i.e.,  in  a 
single  thick  gelatinous  envelope.  Individual  eggs  vary  little  in  size,  each  being 
about  2.3  mm.  in  diameter.  The  gelatinous  capsules  also  are  quite  uniform  at 
about  6.7  mm.  The  gelatinous  coats  are  elastic  and  quite  sSticky  so  that  they 
easily  pick  up  particles  from  the  surrounding  water.  This  tends  to  conceal 
them  after  a  few  hours.  The  little  gelatinous  balls  tend  to  stick  together  in 
various  patterns.  These  patterns  are  quite  irregular,  but  most  often  the  eggs 
attach  tandem-fashion  so  that  an  irregular  string  is  produced  with  some  simi- 
larity to  the  eggs  of  toads.  There  is  no  continuous,  gelatinous,  encasing  tube 
as  in  Bufo,  and  one  egg  within  its  covering  can  easily  be  separated  from  others 


SPADEFOOTS:  SCAPHIOPODIDAE 


133 


to  which  it  is  joined.  Such  strings  may  be  very  short  or  several  inches  long. 
Individual  eggs  of  one  such  string  commonly  attach  to  others  in  adjacent 
strings,  forming  a  network.  Several  such  lattices  are  sometimes  similarly 
joined  to  form  a  three-dimensional  pattern.  A  winding  string  of  eggs,  loosely 
attached  to  other  such  strings  wherever  contact  happens  to  be  made  between 
them,  is  most  commonly  found. 

"The  number  of  eggs  in  a  complete  clutch  is  unknown,  but  the  complement 
must  consist  of  several  hundred  at  least"  (A.  N.  Bragg,  Okla.,  1944,  pp.  232- 


Journal  notes:  On  Feb.  9,  1932,  Mr.  A.  J.  Kirn  of  Somerset,  Tex.,  sent  us  ten 
preserved  Scaphiopus  which  Mr.  Strecker  pronounced  S.  h.  hurterii.  On  Feb. 
15,  1932,  Mr.  Kirn  wrote,  "They  [the  spadefoots]  are  not  calling  as  yet.  I 
found  one  of  them  last  year,  January  27  —  in  the  road,  during  a  rainy  spell. 
I  will  send  you  some  of  the  Scaphiopus  when  they  come  out  to  lay.  Do  not 
know  when  this  will  he."  His  field  notes  for  these  ten  spadefoots  are  some- 
what as  follows:  "Lyile  June  29  (Monday)  1931.  Rain  Friday  8  P.M.  through 
most  of  Saturday  and  again  yesterday,  early  morning  and  late  evening,  again 
early  this  morning  and  this  afternoon,  about  three  inches  altogether.  Weather 
warm.  Scaphiopus  hurterii  heard  last  night  at  all  pools  at  wells  (old  slush 
pits).  Collected  a  dozen  at  No.  10  and  No.  13  (4  pairs  in  copulation;  in  all  of 
these  the  smaller  and  lighter  one  was  male).  Collected  between  10  and  n  P.M., 
June  28.  This  afternoon,  I  found  eggs  already  hatched  at  Nos.  10  and  13  wells. 
No  spadefoots  heard  before  8:30  P.M.  yesterday.  Eggs  evidently  laid  last  night 
after  midnight.  Were  hatched  at  7  P.M.  today  and  how  long  before  I  do  not 
know.  The  spadefoots  collected  last  night  varied  from  gray  and  yellowish 
green  to  dark,  all  with  two  dorsal  stripes  (widening  on  middle  posterior 
back).  Parotoid  glands  distinct  in  all.  Had  in  can  all  of  today.  Many  eggs  laid 
m  the  can.  Not  any  spadefoots  heard  or  found  tonight,  June  29." 

We  had  a  group  of  spadefoots  in  a  laundry  tub,  in  which  we  had  $  inches 
of  dirt.  We  fed  them  mealworms,  from  a  glass  dish  in  the  center  of  the  tub. 
By  clay  no  toads  were  visible,  nor  was  there  any  evidence  of  their  burrows, 
but  about  9:30  in  the  evening  they  began  to  shake  their  heads  and  loose  spots 
appeared  here  and  there,  and  slowly  the  toads  came  out,  and  quickly  snapped 
up  the  proffered  worms. 

We  have  no  personal  experience  with  this  form  in  the  field.  Our  latest  lot 
(1945)  of  live  material  came  from  A.  J.  Kirn,  who  knows  it  best.  Some  of  his 
notes  are: 

"2/12.  Much  rain  last  2  nights.  Spadefoot  toads  calling  loudly  tonight  7:30 
on.  From  water  pools.  This  first  heard  this  spring  and  winter.  Weather  mild. 
2/12.  Very  heavy  rain  last  night  and  nearly  as  much  night  before  that.  Many 
spadcfoot  toads  calling  from  all  pools.  Temperature  57  at  H  P.M.  2/13.  Egg 
clusters  of  spadefoot  toads  this  A.M.  in  a  pool.  Also  numbers  of  tadpoles,  vari- 
ous size  from  small  (%  inch)  to  large,  hind  legs  showing  beneath  skin.  Evi- 


I34  HANDBOOK  OF  FROGS  AND  TOADS 

dently  eggs  laid  some  time  ago.  (January?)  See  2/21.  2/17.  A  few  spadefoots 
in  roads  tonight.  2/21.  Tadpoles,  large,  of  2/13— have  hind  legs  today.  Egg 
cluster  pool  of  2/13  has  numbers  of  tadpoles  about  6  mm.  2/27.  Spadefoot  tad- 
poles of  Feb.  12-13  still  not  as  large  as  the  largest  seen  Feb.  13.  Therefore  those 
were  from  eggs  laid  in  January,  possibly  about  January  21.  This  is  my  first  and 
only  record  of  eggs  laid  during  that  month,  though  I  always  contended  that 
they  would  lay  eggs  any  month  of  the  year  if  temperature  and  conditions  were 
right.  Weather  colder  Feb.  26  and  27.  March  i.  Milder  and  foggy. 

"7/5.  On  June  19, 1  sent  you  a  box  with  25  spadcfoot  frogs,  hope  that  they 
reached  you  alive,  and  that  they  were  enough  to  suit  your  needs.  We  had  a 
four-inch  rain  the  night  before  and  they  called  until  12:15  noon,  then  sun  came 
out,  they  quit  until  after  sundown,  when  some  came  out.  I  went  to  nearby 
ponds  (tanks)  found  them  scarce,  and  wild.  However  I  managed  to  pick  up 
two  dozen,  then  got  one  in  the  house-yard.  Can  get  more  later  on  when  it  rains 
hard  enough,  if  you  need  them." 

Authorities'  corner: 
II.  M.  Smith,  Tex.,  1937,  pp.  104-108. 
V.  M.  Tanner,  Gen.,  1939,  p.  9. 
A.  N.  Bragg,  Okla.,  1942!^,  p.  506. 


FAMILY  BUFONIDAE 

Genus  BUFO  Laurent! 
Maps  8-14 

Colorado  River  Toad,  Giant  Toad,  Girard's  Toad,  Colorado  Toad 

Bufo  alvarius  Girard.  Plate  XXIV;  Map  8. 

Range:  Imperial  Valley,  Calif.,  up  the  Colorado  River  almost  to  Nevada's 
tip,  up  the  Gila  River  almost  to  New  Mexico's  border,  and  south  into  Sonora, 
Mex. 

Habitat:  Lower  Sonoran  life  zone.  Semiaquatic.  In  the  general  locality  of 
large  permanent  streams  or  the  irrigated  portions  of  our  southwestern  desert 
regions. 

Pima  Co.,  Ariz.  "Bufo  alvarius  is  much  commoner  near  the  Steam  Pump 
than  other  species  of  the  genus.  With  a  few  exceptions  it  is  found  only  in  the 
wet  places  around  the  cattle  watering  troughs  of  the  ranches  in  the  mesquite 
association.  The  only  other  place  where  we  found  it  was  in  a  temporary  road- 
side pond  formed  by  a  heavy  rain  where  Scaphiopus  couchii  and  S.  ham- 
mondti  were  breeding.  These,  however,  had  very  probably  come  from  the 
watering  troughs  of  the  ranch  only  a  few  hundred  yards  distant.  While  Bufo 
alvarius  was  found  in  the  pond  with  the  breeding  Scaphiopus,  none  of  the 
former  were  breeding.  One  specimen  was  found  at  dusk  at  the  mouth  of  a 
canyon  on  the  dry  sandy  bed,  and  a  few  were  found  at  night  by  the  roadside. 
Practically  all  were  found  after  dark,  but  we  discovered  that  their  daytime 
hiding  place  was  in  hollows  under  the  watering  troughs"  (A.  I.  Ortenburgcr 
and  R.  D.  Ortcnburger,  Ariz.,  1927,  p.  102). 

Size:  Adults,  3M-7  inches.  Males,  80-156  mm.  Females,  87-178  mm. 

General  appearance:  This  is  a  very  large  grayish  or  brownish-green  toad 
with  smooth,  leathery  skin  and  a  few  scattered  small,  rounded  warts.  The  un- 
der parts  are  light.  The  head  is  broad  and  flat,  marked  by  low,  broad,  crescent- 
shaped  crests  curving  around  the  rear  of  the  eye.  These  are  like  fleshy  folds. 
The  canthus  rostralis  is  marked  by  a  ridge  which  turns  down  in  front  of  the 
eye  as  a  preorbital.  One  to  four  white  warts  are  present  back  of  the  angle  of 
the  mouth.  The  parotoids  are  large,  subreniform  in  shape,  spreading  down- 
ward at  the  shoulder.  There  is  a  large  glandular  wart  on  the  femur  and  a  long 
one  or  several  shorter  ones  on  the  tibia.  These  glands  are  the  conspicuous  mark 


136 


HANDBOOK  OF  FROGS  AND  TOADS 


of  this  toad  and  appear  early,  being  present  in  a  r%-inch  (44-mm.)  individual. 
Color:  Female.  Bard,  Imperial  Co.,  Calif.,  from  L.  M.  Klauber,  May  22, 
1930.  Color  deep  olive,  dark  olive  or  citrine-drab,  or  dark  greenish  olive  on 
back  and  buflfy  olive  on  either  side  of  back.  Warts  on  back  not  conspicuous, 
buckthorn  brown  or  Dresden  brown.  Some  of  same  color  on  each  upper  eye- 
lid. This  brown  scanty  on  femur,  absent  on  forelegs,  tibia,  and  hind  foot.  Fore 
limbs  deep  grayish  olive.  Arm  insertion  olive-gray  and  some  spots  of  same 
color  each  side  of  pectoral  region  and  edge  of  throat.  Throat  same  color  as 
venter.  Venter  white.  Back  of  angle  of  mouth  are  two  pale  ochraccous-buff  or 
shell  pink  glands.  Tympanum  like  dorsal  color.  Iris  cream-buff  below  and 


MapS 

above,  with  some  tawny  or  russet  streaks.  Dark  longitudinal  bar  through  eye, 
brownish  olive  ahead  of  and  behind  eye. 

A  young  specimen  (USNM  no.  21802,  44  mm.)  has  a  spotted  back  and  at 
first  appearance  looks  like  a  B.  punctatits.  The  parotoicls  are  oblong,  spreading 
apart,  the  glands  on  femur  arc  close  to  tibia,  and  those  on  tibia  barely  show. 
Tympanum  very  distinct,  slightly  elliptic.  Two  white  warts  back  of  angle  of 
mouth  present. 

Structure:  Two  metatarsal  tubercles;  two  large  palmar  tubercles;  first  finger 
about  equal  to  second;  first  finger  of  female  may  look  very  long  and  slender, 
that  of  male  much  heavier  at  base;  palms  and  soles  tuberculate;  interorbital 
much  wider  than  internasal  space;  a  membranous  fold  at  the  inner  edge  of 
the  tarsus;  horny  excrescences  on  fingers  of  male  may  be  very  prominent, 
starting  from  back  of  wrist  and  extending  all  along  inner  side  of  first  finger 


TOADS:  BUFONIDAE 


137 


and  covering  upper  surface  as  well;  second  finger  has  upper  surface  with  ex- 
crescence as  well  as  triangular  patch  from  tip  backward;  slight  on  third;  tym- 
panum may  be  obliquely  vertical  and  elliptical  or  almost  round  and  very  little 
oblique. 

Voice:  "When  held  in  the  hand,  this  toad  jerks  spasmodically,  and  vibrates 
the  whole  body,  as  if  about  to  explode  with  wrath.  The  only  sound,  however, 
produced  in  protest  is  a  gentle  chirping  note,  less  loud  and  emphatic  than  that 
of  the  American  toad"  (M.  C.  Dickerson,  Gen.,  1906,  p.  108). 

"I  assure  you  there  was  no  lack  of  noise  that  day  nor  night,  the  croaking 
being  incessant"  (A.  G.  Ruthven,  quoting  J.  J.  Thornber,  Ariz.,  1907,  p.  506). 

"On  the  night  of  July  T3th,  a  chorus  of  giant  toads  was  heard  from  a  ditch 
near  the  Sells-Roblcs  road  southwest  of  Tucson"  (C.  F.  Kauffeld,  Ariz.,  1943, 

P-343)- 
"Of  batrachians,  a  toad  (Bufo)  and  a  frog  (Runa  virescens  brachycephala 

Cope)  were  found  at  Warsaw  Mills;  and  at  Buenos  Ayres,  at  the  beginning 
of  the  summer  rains,  Lieutenant  Gaillard  observed  great  numbers  of  a  very 
large  froglike  toad,  named  Bufo  alvarhts  by  Girarcl.  Nothing  was  seen  or 
heard  of  them  until  the  advent  of  the  early  summer  rains,  which  formed  a 
large  shallow  lake  near  Buenos  Ayres  and  about  10  kilometers  (6  miles)  north 
of  the  Boundary  Line.  These  large  toads  then  filled  the  air  with  their  loud 
cries,  which  increased  until  a  deafening  roar  was  produced.  Numbers  of  them 
were  seen  hopping  about,  but  their  rarity  was  not  suspected  by  Lieutenant 
Gaillard,  on  which  account  none  were  collected"  (E.  A.  Mearns,  Ariz.,  1907, 

P-"3)« 
Breeding:  "Deep  brown  above  and  tan  below,  the  eggs  arc  encased  in  a 

single  long  tube  of  jelly.  The  envelope  is  somewhat  loose,  but  quite  distinct 
in  outline;  the  gelatinous  material  is,  for  the  most  part,  clear,  transparent,  and 
not  very  adhesive.  There  arc  no  partitions  between  the  individual  eggs,  whose 
arrangement  varies  from  a  perfectly  linear  scries  of  near  spheres  to  a  zigzag 
pattern  of  broadly  wedge-shaped  eggs.  The  number  of  eggs  per  inch  averages 
eighteen  (range  12  to  28).  Vitcllinc  membranes  arc  close  to  the  vitclli  and  not 
visible  with  the  unaided  eye;  at  times  they  are  difficult  to  see  even  under  mag- 
nification. Measurements  on  the  eggs  are  as  follows:  vitellus  1.4  mm.  (range 
1.14  to  1.70  mm.) ;  vitellme  capsule  1.6  mm.  (range  1.25  to  i  .70  mm.) ;  envelope 
2.2  mm.  (range 2.12  to  2.25  mm.).  Affinities:  Except  for  the  deeper  color  of  the 
vitelli,  the  lesser  degree  of  convolution  and  lack  of  slight  scalloping  of  the 
gelatinous  tube,  eggs  of  B.  alvartus  are  similar  to  those  of  B.  compactihs.  Meas- 
urements of  both  average  approximately  the  same  m  all  respects.  The  total 
mass  of  eggs  is  larger  in  B.  alvariits  and  the  crowding  within  the  tube  is  more 
intense  (12  to  28  per  inch  for  alvarius  to  14  to  20  for  compactilis),  this  differ- 
ence, however,  being  barely  noticeable.  Superficially  B.  alvarius  eggs  also  look 
like  those  of  B.  terrestris.  But  here  the  lighter  color  of  the  eggs  and  greater 


138  HANDBOOK  OF  FROGS  AND  TOADS 

degree  of  convolution  of  the  tube  plus  the  presence  of  an  inner  envelope  serve 
to  separate  eggs  of  these  species  immediately."  (R.  Livezey  and  A.  H.  Wright, 
Gen.,  1947,  pp.  193, 194,214). 

The  eggs  we  secured  July  2-3,  1934,  were  *n  l°n&  ropelikc  strings  (400 
inches)  of  7500-8000  eggs. 

"Two  of  the  females  collected  at  Alamos  between  August  27  and  Septem- 
ber 2  contain  mature  eggs  in  the  oviduct"  (C,  M.  Bogert  and  J.  A.  Oliver, 
Gen.,  1945,  p.  339). 

Journal  notes:  July  30,  1917,  just  southeast  of  Tempe,  Anz.  Made  only  49 
miles.  In  a  water  hole  near  a  culvert  Ralph  Wheeler  and  I  caught  six  immense 
toads  (Bufo  alvanus).  All  males.  Probably  tardy  ones.  Tadpoles  (Bufo)  in  the 
hole  probably  of  this  species. 

June  25, 1934,  Florida  Experiment  Station,  Ariz.  Mr.  R.  R.  Humphrey  said 
that  at  10  P.M.  on  the  road  from  Nogales  north  he  saw  several  Bufo  alvanus 
after  rain.  Mr.  Gorsuch  said  they  often  gather  at  the  Continental  pond,  that  he 
once  found  several  pairs  of  B.  alvanus  which  he  brought  up  to  the  station. 
They  laid  long  strings  of  black  eggs.  Had  he  known  they  were  not  described 
he  would  have  saved  them. 

June  29.  At  Continental  we  walked  around  an  irrigation  pond.  Under  a  tin 
cover  found  a  large  male  B.  alvanus.  How  he  did  protestingly  chuckle  as  we 
carried  him  off.  His  body  under  my  hand  was  clammy  with  secretions. 

July  2,  Tucson,  Ari/.  Rained  in  many  places.  Went  about  5  to  Sabmo  Can- 
yon. At  picnic  spot  two  pools,  the  upper  pool  filled  with  R.  pipiens.  When  al- 
most dark  we  went  to  the  lower  pool  with  its  dam.  At  upper  end  we  saw  about 
12  dead  females  of  Bit  jo  alvarius.  Several  large  B.  alvanus  males  jumped  in. 
We  continued  to  find  males  until  we  came  to  where  Prof.  Wehrle's  small 
daughter  had  spotted  a  just  perceptible  gray  spot  in  the  shallow  water  (4-8 
inches).  This  proved  to  be  a  pair.  There  were  six  or  eight  pairs  here.  Brought 
in  three  pairs  and  put  them  in  a  tub.  How  the  males  did  cluck!  They  sound 
like  contented  chickens  or  chickens  when  sleepy  at  night.  At  ti  P.M.  we  found 
they  had  laid  and  we  fixed  some.  What  immense  ropelike  masses!  They 
clucked  when  disturbed. 

July  3.  This  morning  many  B.  alvanus  eggs  in  the  tub.  One  pair  yet  mated. 
The  other  pairs  are  broken.  Went  to  Sabino  Canyon.  No  B.  alvanus  to  be 
found  or  eggs. 

Authorities'  corner: 

J.  G.  Cooper,  Calif.,  1869,  p.  480.  A.  I.  and  R.  D.  Oitenburger,  Ariz.,  1927, 

F.  Mocquard,  Gen.,  1899,  p.  168.  p.  102. 

E.  A.  Mearns,  Ariz.,  1907,  p.  113.  F.  W.  King,  Ariz.,  1932,  p.  175. 

A.  G.  Ruthven,  Ariz.,  1907,  p.  506.  C.  F.  Kauflfeld,  Ariz.,  1943,  p.  343. 


TOADS:  BUFONIDAE 


'39 


Plate  XXIV.  Bufo  alvarius.  1,2,3,4.  Fe- 
lales 


Plate  XXV.  Btifo  americanus  amcncanus. 
1,2,3,7.  Males  (X1/^)-  4-  Male  trilling  while 
sitting  in  shallow  water  (X%).  5-  YounS 

(X1'-')-  6-  Coils  of  e^  strinSs  on  bottom 
of  a  pond 


I40 


HANDBOOK  OF  FROGS  AND  TOADS 


American  Toad,  Northern  Toad,  Hop  Toad 

Bujo  amencanus  amcncanus  Holbrook.  Plate  XXV;  Maps  9,  10. 

Range:  Manitoba  eastward  on  latitude  50°  N.  to  Gaspe  and  the  Maritime 
Provinces.  Minnesota  south  through  Iowa  to  eastern  Kansas  and  Oklahoma, 
thence  eastward  to  northern  Georgia,  along  Appalachians  to  central  Virginia, 
thence  to  the  coast  and  northward  to  Nova  Scotia.  A  transition  and  Canadian 
zone  form.  Common  throughout  its  range. 

Habitat:  Common  in  gardens  and  cultivated  fields,  appearing  more  by 
night  than  day.  During  the  sunshiny  hours  they  seek  cover  beneath  piazzas, 


B.  a.  cope? 


Map  9 


TOADS:  BUFONIDAE 


141 


under  board  walks,  flat  stones,  boards,  logs,  wood  piles,  or  other  cover.  When 
cold  weather  comes,  the  toad  digs  backwards  into  its  summer  quarters  or  may 
choose  another  site  for  its  hibernation. 

Size:  Adults,  2^-4 Vi  inches.  Males,  54-85  mm.  Females,  56-110  mm. 

General  appearance:  Short  and  fat  in  body,  it  has  a  short  broad  head  and 
the  snout  is  broadly  circular.  The  lower  surfaces  are  roughly  granular,  the 
back  is  covered  with  various-sized  warts,  some  of  which  are  large  ones  in  pairs 
down  the  middle  of  the  back.  There  arc  three  or  four  pairs  of  dark  spots  down 
the  back,  each  with  one  large  wart.  The  eyes  are  prominent.  The  arms  and 
legs,  hands  and  feet,  are  warty  or  roughly  tubercular.  There  arc  dark  spots  on 
the  arms  and  legs,  along  the  sides,  and  a  few  on  the  belly.  Some  males  have 
yellow  throats  and  considerable  yellow  on  the  underside  of  the  base  of  the  legs 
and  in  the  groin.  The  general  color  is  olive,  with  parotoids  and  crest  brown. 

Color:  Male.  May  7,  1929.  Back,  sides,  and  tympanum  dull  citrine  with 
olive-citrine  or  yellowish  olive  on  hind  legs  and  forelegs.  Parotoids,  crests, 
snout,  and  face  ahead  of  eye  huffy  olive.  Throat  old  gold,  olivc-ocher,  aniline 
yellow,  or  yellow  ocher.  Some  of  same  color  in  axilla  and  on  rear  of  brachium, 
also  on  groin  and  on  lower  belly.  In  latter  region  may  be  ochraccous-buff. 
Whole  pectoral  region  with  distinct  scattered  black  spots;  in  fact  these  occur 
over  entire  venter  except  throat  and  center  of  rear  belly.  Lower  throat  oil- 
yellow  or  sulphine  yellow.  Some  apricot  yellow  across  arm  insertion.  Sides 
heavily  spotted  with  large  spots  of  black.  On  back  is  an  obscure  oblique  black 
bar  bordered  with  greenish  yellow  on  upper  eyelid.  It  is  interrupted  by  the 
superciliary  crest,  only  a  small  part  showing  inside  this  structure.  One  buffy 
olive  tubercle  in  center  of  the  eyelid  is  almost  on  the  outer  end  of  this  bar.  A 
more  or  less  round  black  spot,  greenish-yellow-edgcd,  is  just  inside  front  end 
of  long  parotoid  and  just  back  of  postorbital  bar  and  end  of  superciliary  crest, 
with  tubercle  in  the  center.  Opposite  the  rear  portion  of  parotoid  are  two 
oblique,  posteriorly  divaricating  narrow  bars  with  tubercles  on  forward  end 
of  each.  Just  beyond  end  of  parotoids  are  the  two  largest  round  black  spots 
with  a  large  tubercle  in  each  center.  Halfway  from  parotoid  to  vent  another 
such  pair.  Back  of  this  pair  arc  suggestions  of  two  or  three  obscure  pairs.  The 
tubercles  in  the  five  pairs  are  buffy  olive.  Femur  and  tibia  are  more  or  less 
crossbarrcd  with  black.  Fore  limbs  spotted  with  black  and  a  prominent 
oblique  black  spot  across  front  of  arm  insertion.  Rear  of  femur  spotted  or 
vermiculated  with  black  and  olive-citrine,  old  gold,  olive-ocher,  or  primulinc 
yellow.  Iris:  dark  in  front  of  and  behind  pupil.  Pupil  rim  citron  yellow.  Above 
pupil,  mustard  yellow  and  ochraceous-buff. 

Female.  July  24, 1929.  Color  of  back,  light  brownish  or  buffy  olive;  parotoids 
isabella  or  dark  olive-buff;  the  bigger  warts  on  the  back,  which  arc  in  centers 
of  dark  spots,  buffy  brown;  crests  also  buffy  brown;  stripe  down  middle  of 
back,  deep  olive-buff,  yellowish  glaucous,  or  pale  vinaceous-fawn,  with  more 
or  less  of  the  same  color  on  oblique  stripe  that  leads  from  parotoid  to  groin, 


142  HANDBOOK  OF  FROGS  AND  TOADS 

and  also  along  the  sides  above  and  below  the  dark  lateral  band.  The  dark 
lateral  band  is  brownish  olive,  olive,  or  deep  olive.  The  tympanum  may  be 
buff  olive  or  light  grayish  olive.  The  paired  spots  down  the  middle  of  the  back 
are  (i)  oblique  bars  on  upper  eyelids;  (2)  spots  inside  the  front  of  parotoid; 
(3)  another  pair  of  about  the  same  size  at  rear  end  of  parotoid  with  a  spot 
near  the  middle  line  between  these  two.  Back  of  these  and  farther  from  the 
middle  line  are  (4)  two  larger,  usually  two-tuberclcd,  spots  sometimes  with 
two  small,  single-tubcrcled  spots  inside  each  of  these  and  nearer  the  meson. 
The  next  pair  (5)  are  about  the  middle  of  the  back  and  close  to  the  meson. 
These  arc  the  most  prominent  of  the  back  and  arc  from  i-  to  4-tuberclcd.  Then 
back  some  distance,  close  to  the  meson  and  slightly  ahead  of  the  vent,  is  an- 
other pair  (6).  These  spots  are  black,  fuscous  black,  or  clove  brown  and  are 
surrounded  by  reed  yellow  borders.  Below  the  eye  and  tympanum  is  a  black 
spot  with  a  snuff  brown  center.  The  side  of  the  face  is  clay  color  or  huffy  pink. 
The  iris  is  same  as  in  the  male.  Under  parts  are  cream  buff  on  the  throat,  be- 
coming primrose  yellow  on  the  belly  and  honey  yellow  or  deep  colonial  buff 
on  middle  of  lower  belly  and  inner  portion  of  buttocks.  Among  this  is  some 
orange-vmaceous  or  pale  grayish  vinaceous.  In  center  of  breast  is  a  dark  spot. 

Structure:  Pnrotoids  large  and  oblong,  connected  to  the  postorbital  crest  by 
a  longitudinal  ridge.  Crests  on  the  head  form  a  right  angle  at  the  corner  of 
the  eye,  one  branch  extending  downward  in  front  of  the  ear;  males  with  dark 
throats;  males  with  excrescences  on  inner-upper  side  of  first  two  fingers  and 
on  inner  carpal  tubercle;  many  spiny  warts,  particularly  on  the  hind  limbs. 

Voice:  "The  note  of  the  toad  is  long  sustained,  quite  musical,  rather  high 
in  pitch.  Their  low  variable  trills  or  pipings  in  chorus  are  rather  pleasing  and 
do  not  especially  annoy,  even  though  continued  day  and  night  during  the 
height  of  their  breeding  season"  (A.  H.  Wright,  N.Y.,  1914,  p.  27). 

"In  general  it  sounds  like  whistling,  and  at  the  same  time  pronouncing  deep 
in  the  throat,  bu-rr-r-r-r.  It  will  be  found  that  different  toads  have  slightly  dif- 
ferent voices,  and  the  same  one  can  vary  the  tone  considerably,  so  that  it  is 
not  so  easy  after  all  to  distinguish  the  many  batrachian  solos  and  choruses  on 
a  spring  or  summer  evening"  (S.  H.  Gage,  N.Y.,  1904,  p.  186). 

"The  common  toad  of  the  mainland  of  New  York  State  is  called  Bttfo  amer- 
icanns.  Its  song  is  a  sweet,  trilling  whistle,  and  may  be  imitated  by  whistling 
in  a  low  monotone  with  drops  of  water  held  between  the  lips.  Each  individual 
song  is  prolonged  for  about  thirty  seconds.  The  prolonged  song  of  the  Ameri- 
can toad  is  a  ready  means  of  distinguishing  it  from  the  short  song  of  the 
common  toad  (jowleri)  of  Long  Island"  (F.  Overton,  N.Y.,  1914,  p.  27). 

Breeding:  They  breed  from  April  5  to  July  25,  the  crest  about  April  30.  The 
eggs  are  in  long  spiral  tubes  of  jelly,  each  egg  %s-Me  inch  (1.0-1.4  mm.)  in 
diameter;  the  inner  tube  %-]i2  inch  (1.6-2.2  mm.),  the  outer  tube  %-% 
inch  (3.4-4.0  mm.).  The  eggs,  4000-8000  in  number,  are  laid  in  two  strings, 
and  hatch  in  3-12  days.  The  small,  dark,  almost  black  tadpole  i%2  inches  (27 


TOADS:  BUFONIDAF  143 

mm.)  has  an  ovoid  body  broader  near  the  vent  than  at  the  eyes.  The  dorsal 
crest  is  low,  extending  slightly  onto  the  body,  the  tail  short,  its  tip  rounded. 
The  tooth  ridges  are  73.  After  50-65  days,  the  tadpoles  transform  June  i  to 
August  at  /4-Vis  inch  (7-12  mm.). 

Journal  notes:  June  8, 1911.  Found  at  Crossroads  Pond,  Ithaca,  N.Y.,  numer- 
ous Bufo  americanus  transformed  and  transforming,  also  several  Rana  syl- 
vatica  transformed  (at  least  12  or  15).  At  Beebe  Lake  on  south  side  near  the 
spring  are  myriads  of  transformed  Bujo  americanus. 

June  9,  Beebe,  north  side.  On  this  side  also  arc  countless  Bujo  americanus 
transformed.  John  Rich  reports  them  numerous  yesterday  on  Wait  Avc.  just 
over  Triphammer  bridge.  Beebe,  south  side,  Bufo  transformed  still  numerous. 

June  12,  Beebe,  by  the  spring.  Little  toads  are  crossing  the  road  above.  They 
are  not  so  numerous  at  water's  edge. 

April  23,  1912,  Crossroads  Pond.  Found  two  pairs  of  Bufo  americanus  lay- 
ing. Water  cold,  not  much  warmer  than  4<)°F.  Found  hidden  males  at  varying 
distances  from  the  pond  on  the  hillside  below.  Many  of  them  were  not  entirely 
under  cover.  They  were  in  little  pockets  at  the  surface  of  the  ground,  their 
backs  exposed  and  the  skin  as  dark  and  dirtlikc  as  the  dirt  itself. 

Authorities9  corner: 
C.  C.  Abbott,  N.J.,  1868,  p.  805. 
A.  II.  Kirkland,  Mass.,  1897,  p.  24. 
H.  Carman,  Ky.,  1901,  pp.  60-61. 

Hudson  Bay  Toad,  Hudson  Bay  American  Toad,  Cope's  Toad 

Bufo  americanus  copei  Yarrow  and  Hcnshaw.  Plate  XXVI;  Maps  9, 10. 

Range:  Hudson  Bay,  James  Bay.  Gaige  (Out.,  1952,  p.  134),  who  brought  it 
out  of  synonymy,  wrote:  "During  a  vacation  trip  to  the  James  Bay  region  in 
August  of  this  year  Mr.  Calvin  Goodrich  kindly  collected  a  scries  of  toads  ( 10 
.specimens,  ranging  from  10  mm.  to  69  mm.  in  length)  for  us  at  Moose  Fac- 
tory." Riviere  clcs  Rapides,  Labrador  (F.  G.  Speck,  Lab.,  1925,  pp.  5-6); 
Quebec. 

Habitat:  "The  colony  .  .  .  inhabits  the  abandoned  clearing  of  a  lumber 
operation  of  some  years  ago.  Three  or  four  decayed  log  stables  and  sheds  and 
a  surrounding  mass  of  fine  succulent  meadow  grass  near  the  mouth  of  the 
river  formed  the  environment.  This  small  race  is  very  active,  and  I  have  no- 
ticed that  the  living  specimen  spends  much  time  buried  about  an  inch  deep  in 
the  sand  .  .  ."  (F.  G.  Speck,  Lab.,  1925,  pp.  5-6). 

Size:  Adults  1%-j  inches  (40-75  mm.). 

General  appearance:  "The  brilliant  coloration,  the  long,  narrow  parotoid 
glands,  the  greater  width  between  the  cranial  crests  and  the  fact  that  they  are 
nearly  parallel,  are  the  most  obvious  characters.  The  smoothness  of  the  ventral 
granulation,  mentioned  by  Henshaw  and  Yarrow,  is  evident,  but  it  may  be 


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TOADS:  BUFONIDAE  145 

seasonal  or  due  to  preservation,  and  the  hind  limbs  are  shorter  than  is  usual 
in  Michigan  specimens  of  B.  amcricanns,  the  adpressed  heel  reaching  the 
shoulder"  (H.  T.  Gaige,  Ont.,  1952,  p.  134). 

Color:  Thirty  miles  cast  of  Moisie,  Labrador,  from  H.  W.  Jackson,  1937. 
In  captivity  the  male  lived  a  year,  the  female  two  years. 

Female.  The  larger,  more  brightly  colored  of  the  two.  The  groin  and  rear 
axilla  are  brazil  red.  There  is  also  a  wash  of  the  same  color  on  rear  under  parts. 
A  few  dorsal  tubercles  along  mid-back  are  claret  brown  or  chestnut  brown. 
The  snout  and  parotoid  glands  are  hazel.  The  middorsal  stripe,  light  lateral 
stripe,  and  other  light  dorsal  spots  are  pale  smoke  gray.  The  under  surface  is 
cartridge  buff  with  heavy  black  mottling.  The  iris  has  glass  green  pupil  rim 
and  flecks  of  the  same  color. 

Male.  The  smaller  toad  is  almost  completely  uniform  in  color.  The  snout, 
crests,  parotoid  glands,  and  warts  are  isabclla  color.  The  back  is  olive- 
ocher  or  ecru-olive.  The  spots  on  rear  of  dorsum  are  yellowish  olive,  a  few 
near  the  head  are  black.  The  belly  is  pale  olive-buff,  heavily  spotted  with  black. 
The  groin,  rear  of  femur,  and  to  a  slight  extent  axilla  are  apricot  orange.  In 
the  eye,  the  pupil  rim  is  light  orange-yellow  with  heavy  flecking  of  the  same 
in  upper  part  of  the  iris. 

Structure:  "Head  subtriangular,  broader  than  long;  snout  acuminate,  pro- 
truding; head  with  well-marked  groove,  which  extends  to  tip  of  snout;  super- 
ciliary ridges  strongly  pronounced  and  terminating  posteriorly  in  a  slight 
knob;  orbit  bordered  posteriorly  by  a  similar  ridge;  upper  jaw  slightly  emar- 
ginatcd;  parotoids  medium,  elongated,  twice  as  long  as  broad,  perforated  by 
numerous  small  pores,  situated  well  back  on  the  shoulders;  not  approximated 
to  the  tympanum,  which  is  circular  and  large;  limbs  long  and  comparatively 
slender;  palm  rugose;  a  single  well-developed  tubercle;  first,  second,  and 
fourth  fingers  about  equal  in  length,  the  third  longest;  hind  limbs  rather 
longer  than  head  and  body  together;  tarsus  and  metatarsus  with  small  and 
smooth  tubercles;  body  above  covered  with  small  and  somewhat  roughened 
tubercles;  under  parts  finely  papillatcd;  metatarsal  shovel  large"  (H.  C.  Yar- 
row and  H.  W.  Hcnshaw,  Ncv.,  1878,  p.  207). 

Voice:  Not  of  record. 

Breeding:  Not  of  record. 

Journal  notes:  We  have  made  notes  on  the  Clausen,  Trapido,  and  Vladykov 
material.  Several  explorers  in  Gaspe,  Labrador,  and  Hudson  Bay  have  re- 
marked :  "What  is  the  bright  tood  we  saw  ?" 

Authorities9  corner:  "Toads  common  in  the  valley  of  the  Ste.  Anne  des 
Monts  River,  Gaspe  Peninsula,  differ  strikingly  from  Bufo  a.  americanus. 
Comparison  at  the  Canadian  National  Museum  with  material  of  B.  ameri- 
canus copei  from  Moose  Factory  and  from  Cape  Hope  Islands  in  James  Bay 
demonstrated  that  our  toads  were  that  subspecies.  The  coloration  is  much 
brighter  than  in  B.  a.  americanus,  with  the  black  spots  of  the  back  fused  on 


146  HANDBOOK  OF  FROGS  AND  TOADS 

the  sides  to  form  broken  bands  and  blotches.  These  markings  are  usually  so 
arranged  that  there  is  a  broad  light  middorsal  band,  and  frequently  a  light 
lateral  stripe  on  each  side,  from  the  parotoid  to  the  insertion  of  the  hind  limb. 
Our  specimens  also  resemble  B.  a.  copei  in  the  pink  coloration  in  the  axils  of 
the  limbs  and  in  the  pink  tipped  warts  over  the  dorsal  body  surface,  as  Cope 
(1895:  286)  noted  for  a  specimen  from  Moose  River,  British  America,  which 
he  assigned  to  copei  of  Yarrow  and  Henshaw,  although  he  did  not  consider 
that  form  valid.  The  parotoids  not  only  are  long  and  narrow,  as  Gaige  (1932) 
has  pointed  out,  but  tend  to  be  constricted  towards  their  middle,  while  the 
chest  and  belly  are  speckled  or  in  some  individuals  strongly  mottled  with 
black. 

"Sexual  dimorphism  is  pronounced  in  a  pair  of  living  specimens  which  we 
have  examined  from  Seal  River,  on  the  north  shore  of  the  St.  Lawrence  River. 
The  female  is  brilliantly  colored  with  black  and  pink  dorsally,  and  with  black 
marbling  vcntrally,  while  the  male  is  a  drab  yellow  brown,  with  the  black  and 
pink  much  less  pronounced  and  the  middorsal  stripe  lacking.  The  underside 
of  the  male  is  spotted  with  black,  though  not  so  extensively  as  in  the  female. 
The  parotoids  of  both  male  and  female  are  narrow  and  long,  while  the  cranial 
crests  are  well  spaced.  Our  collection  from  the  Ste.  Anne  River  differ  from  this 
pair  in  that  the  males  arc  as  brilliantly  colored  as  the  females"  (H.  Trapido 
and  R.  T.  Clausen,  Que.,  1938,  p.  120). 

"Bttfo  americanus  americanus  (Holbrook).  Common  throughout  the  area 
and  very  abundant  locally;  usually  found  on  land,  but  a  few  taken  from  wa- 
ter. Most  of  the  toads  were  brilliantly  colored,  with  patches  of  pink  below  and 
many  small  pink  tipped  warts  on  the  sides.  The  dorsal  spots  were  deep  black, 
with  several  large  warts  in  each.  These  toads  thus  tend  to  resemble  B.  ameri- 
canus copei  Yarrow  and  Henshaw  in  coloration  but  lack  the  structural  pecu- 
liarities of  that  subspecies.  Logier  (1928)  has  noted  the  more  brilliant  colora- 
tion of  northern  toads.  The  call  was  heard  occasionally  until  July  17" 
(R.  Grant,  Que.,  1941,  p.  151). 

Northwestern  Toad,  Baird's  Toad,  Mountain  Toad,  Columbian  Toad,  Small- 
spaded  Toad,  Northern  Toad,  Western  Toad 

Btifo  boreas  boreas  (Baird  and  Girarcl).  Plate  XXVII;  Maps  10,  n. 

Range:  Prince  William  Sound,  southeastern  Alaska,  southeastward  to  east- 
ern Montana  and  northeastern  Colorado  and  southwestern  Utah,  thence  west 
to  California  (Mono  County  to  northwestern  California). 

Habitat:  Terrestrial  except  at  breeding  time.  Ruthven  and  Gaige  (1915) 
stated  the  situation  very  well  and  tersely  in  their  table  of  habitat  distribution 
in  Maggie  Basin  in  northeastern  Nevada.  Bufo  b.  boreas  occur  in  "general 
vicinity  of  larger  water  courses.  Breed  in  water  but  can  endure  rather  dry 
habitats  during  other  times  of  the  year."  It  was  common  in  "basin  floor  valleys 


TOADS:  BUFONIDAE 


Plate   XXVI.    Bufo    amerlcanus   copei 
-  1,2,3.  Females.  4.  Male. 


Plate  XXVII.  Bufo  boreas  boreas.  1,2. 
Males  (X1/2).  3.  Hind  foot  of  male  (X%). 
4,5.  Females  ( 


148  HANDBOOK  OF  FROGS  AND  TOADS 

and  mountain  canyons  nearly  to  highest  elevations.  .  .  .  Lives  in  tules  about 
lake  shores,  along  streams,  and  in  mountain  meadows"  (J.  Grinnell  and  C.  L. 
Camp,  Calif.,  1917,  p.  143). 

Size:  Adults,  2^-5  inches.  Males,  56-108  mm.  Females,  60-125  mm. 

General  appearance:  This  is  a  large  brown,  gray,  or  green  toad  with  a  light 
streak  down  the  speckled  back.  The  head  is  narrow  and  pointed  in  proportion 
to  the  fat  broad  body.  There  are  rounded  glands  on  the  middle  of  the  tibia. 
Lack  of  a  discolored  throat  makes  the  males  hard  to  distinguish  from  the 
females. 

Color:  Aug.  31, 1929.  Male.  The  light  middorsal  stripe  is  pale  cendre  green, 
pale  dull  green-yellow,  or  pale  Veronese  green,  or  even  cartridge  buff  pr  sea- 
foam  yellow.  The  ground  color  along  the  back  is  dark  olive  or  clove  brown, 
becoming  about  one-half  inch  from  middorsal  line  pale  brownish  drab,  light 
drab,  or  drab-gray.  As  it  approaches  the  belly  it  becomes  an  ecru-drab  or  pale 
drab-gray.  The  belly  color  is  tilleul  buff  to  pale  vinaceous-fawn,  or  pale  smoke 
gray.  The  throat,  clear  of  spots,  may  be  vinaceous-buff,  tilleul  buff,  olive-buff, 
or  pale  smoke  gray.  Along  either  side  of  the  middorsal  line  is  an  irregular  row 
of  warts.  The  next  series  outside  begin  to  be  conspicuously  surrounded  by 
black  or  black  interspersed  with  citrine-drab  areas.  On  the  lower  side  the  con- 
spicuous warts  in  the  middle  of  the  dark  areas  are  lacking,  and  only  the  citrine- 
drab  or  water  green  spots  are  present.  Across  the  belly  these  large  black  spots 
are  very  much  in  evidence  as  also  on  the  inside  of  the  hind  legs  out  to  the  toes. 
The  underside  of  the  forelegs  also  have  them,  but  they  are  localized  on  the 
rear  and  upper  margins.  On  the  rear  of  the  belly  and  the  underside  of  the 
buttocks  the  black  spots  are  very  fine,  interspersed  with  cream  color  or  light 
vinaceous-fawn.  Here  the  tubercles  are  much  larger  and  mainly  cartridge  buff 
or  pinkish  buff.  The  tubercles  on  the  underside  of  the  fore  feet  and  tips  of  toes 
are  light  vinaceous-cinnamon.  The  parotoid  is  buffy  olive  or  buffy  brown,  as 
is  also  the  rim  of  the  upper  eyelid.  The  upper  jaw  and  face  ahead  of  the  eye  arc 
uniform  grayish  olive.  There  is  a  dark  oblique  area  from  the  lower  rear  of  the 
eye  past  the  angle  of  the  mouth.  There  is  another  dark  area  from  the  middle 
of  the  rear  of  the  eye  over  the  small  tympanum  and  along  the  lower  margin 
of  the  parotoid.  There  is  a  long  longitudinal  bar  from  the  shoulder  insertion 
along  the  front  of  the  forearm.  The  eye  is  black  or  raw  umber  through  the 
middle.  The  rim  of  the  pupil  is  maize  yellow.  The  lower  pupil  rim  is  of  the 
same  color  or  warm  buff,  but  it  is  narrow  and  broken  in  its  middle  point. 
Above  the  upper  pupil  rim  is  the  most  conspicuous  feature  of  the  eye — a  prom- 
inent zinc  orange  or  mikado  orange  longitudinal  band.  The  lower  part  of  iris 
is  dotted  with  ochraceous-buff  or  zinc  orange. 

Structure:  No  cranial  crests;  rounded  gland  in  middle  of  tibia;  body  flat, 
broad;  head  narrow  and  pointed  in  proportion;  no  discolored  throat  in  male; 
spread  of  hind  foot  usually  more  than  36  per  cent  of  total  body  length.  "Warts 


TOADS:  BUFONIDAE  149 

on  back  show  a  tendency  to  run  in  longitudinal  rows.  Tibia  with  one  large 
and  one  small  parotoidlike  wart  located  respectively  in  the  central  and  the 
rear  cross  bars"  (C.  L.  Patch,  B.C.,  1922,  p.  77). 

Voice:  Cope  heard  this  form  in  a  pond  near  the  shore  of  Pyramid  Lake, 
Nev.  It  was  associated  with  a  spadefoot  (Scaphiopus  intcrmontanus)  of  which 
Mr.  Cope  wrote:  "Like  other  allied  species,  it  was  very  noisy,  almost  obscuring 
the  voice  of  the  less  vociferous  Btifo"  (E.  D.  Cope,  Nev.,  1884,  p.  18). 

Our  captive  male  toads,  when  held  in  hand  and  frequently  at  other  times, 
give  little  birdlike  chirps,  very  pleasant  in  tone. 

"At  this  time  the  males  call  with  a  high-pitched  tremulous  note,  amplified 
by  the  vocal  sacs  distended  beneath  the  chin"  (G.  C.  Carl,  B.C.,  194?,  p.  43). 

"The  adults  were  large  and  tame,  they  usually  walked  instead  of  hopped, 
and  when  confined  in  a  bag  they  scolded  much  like  B.  americanus"  (A.  G. 
Ruthven  and  H.  T.  Gaige,  Nev.,  1915,  p.  14). 

Breeding:  They  breed  from  March  to  July.  The  eggs  are  in  strings  like  those 
of  B.  boreas  halophtlits  of  California.  Several  workers  have  seen  the  eggs  but 
none  has  described  them  in  detail  (except  Slater  in  manuscript).  Our  evidence 
is  not  sufficient,  but  it  appears  that  the  eggs  are  as  follows:  Two  envelopes; 
no  partitions  between  the  individual  eggs;  eggs  largely  in  a  double  row  within 
the  jelly  tube;  outer  envelope  large;  4.8-5.3  mm.,  some  loose  and  distinct; 
inner  envelope  3.5-3.8,  distinct;  vitellus  1.5-1.75  mm. 

The  tadpoles  are  small,  i  Vi2  inches  (27-44  mm«)  with  teeth  %.  There  is  no 
description  of  them  on  record.  After  a  tadpole  period  of  30  to  45  days,  they 
transform  from  July  to  September  at  %-Yi  inch  (9.5-12  mm.). 

"On  the  nights  of  June  n  and  12,  1928,  these  toads  were  seen  in  numbers 
in  a  larger  pond  on  the  beach  at  Kaslo  [B.C. |.  The  males  were  calling  and 
greatly  outnumbered  the  females;  nearly  all  the  specimens  collected  or  exam- 
ined at  the  pond  were  males.  One  male  was  seen  on  the  beach  in  embrace  with 
a  dead  female  which  was  much  dried  and  shrivelled.  .  .  . 

"A  female  of  108  mm.  taken  at  Kaslo  on  June  n,  1928,  had  apparently  fin- 
ished spawning;  two  other  specimens  of  81  and  101  mm.  taken  at  Summcrland 
in  July,  one  on  the  i7th,  were  full  of  eggs.  A  specimen  of  93  mm.  taken  at  Lyt- 
ton  between  the  ist  and  8th  of  July,  1925,  had  evidently  spawned. 

"On  the  night  of  June  12,  strings  of  eggs  were  found  strewn  among  the 
grasses  in  the  pond  of  water  six  or  eight  inches  deep.  The  water  temperature 
was  66°  F."  (E.  B.  S.  Logier,  B.C.,  1932,  p.  321). 

"Btifo  boreas  in  Alaska:  In  the  winter  of  1896,  Mr.  A.  W.  Greeley,  a  student 
at  Leland  Stanford  Junior  University,  gave  me  for  examination  2  toads  which 
he  had  'taken  swimming  in  a  large  lake  near  Prince  William's  Sound,  Alaska, 
July  15,  1896.'  .  .  .  One  of  these  specimens  contains  eggs  which  must  have 
been  nearly  ready  for  laying"  (J.  Van  Denburgh,  Alas.,  1898,  p.  139). 

Several  workers  in  Oregon,  Utah,  and  Washington  have  found  this  form 


150  HANDBOOK  OF  FROGS  AND  TOADS 

breeding.  James  Slater  has  sent  us  some  fine  tadpole  material  but  we  are  await- 
ing his  Amphibia  of  Washington  before  venturing  on  further  allusion  to 
them. 

We  have  found  on  record  no  detailed  description  of  the  eggs  or  tadpoles. 

Journal  notes:  March  30,  1942.  Went  with  James  Slater  to  Sparaway  Lake, 
Wash.  As  we  approached  the  lake  we  saw  a  Bufo  boreas  borcas  male.  Caught 
it. 

April  7.  East  of  Ashland  and  where  road  crosses  creek  (junction  with  Samp- 
son Creek)  saw  Bufo  boreas  boreas  run  over. 

April  8.  Up  creek  above  Trail,  Ore.  In  one  pool  took  a  female  and  a  male 
Bufo  boreas  boreas. 

Authorities'  corner: 

E.  D.  Cope,  Ore.,  1883,  p.  19.  C.  E.  Burt,  Gen.,  19333,  p.  351. 

M.  M.  Ellis  and  J.  Henderson,  Colo.,      C.  F.  Schonberger,  Ore.,  1945,  p.  121. 

1915,  p.  254. 

California  Toad,  Salt-Marsh  Frog,  Baird's  Toad,  Common  Toad 

Bujo  boreas  halophilus  (Baird  and  Girard).  Plate  XXVIII;  Map  11. 

Range:  According  to  Camp,  the  seven  northern  counties  of  California  have 
Bufo  b.  boreas,  while  south  of  Eureka  toads  are  Bufo  b.  halophilits.  This  spe- 
cies extends  the  length  of  the  Great  Valley  and  along  the  coast  to  northern 
Lower  California.  Somewhat  east  of  Los  Angeles,  avoiding  the  southeastern 
deserts,  the  range  cuts  diagonally  north  of  Owen's  Lake  to  Lake  Tahoc  or 
Pyramid  Lake  in  Nevada. 

Habitat:  Open  valleys,  rarely  wooded  areas.  In  high  mountains  found  in 
wet  meadows  and  along  lake  shores. 

Size:  Adults,  2?  3-4%  inches.  Males,  62-101  mm.  Females,  fx>-ii6  mm. 

General  appearance:  This  is  a  large  stocky  toad  with  short  limbs,  green  or 
greenish  brown  with  a  light  streak  down  the  back,  and  the  back  mottled  with 
irregular  dark  areas  which  surround  the  warts  singly  or  in  irregular  groups. 
The  warts  are  rounded  and  the  skin  between  the  warts  quite  smooth.  The 
white  or  yellowish  under  parts  are  in  some  individuals  blotched  with  black. 

Color:  April  22, 1928.  Male.  Stripe  down  the  back  is  light  lumiere  green,  or 
chrysolite  green,  or  glass  green.  Upper  parts  mignonette  green,  light  yellow- 
ish olive,  or  yellowish  olive,  becoming  on  the  sides  of  the  head  buffy  olive. 
Yew  green  or  jade  green  on  sides  of  body  with  background  pale  glass  green  or 
glass  green.  Under  parts  white.  Buttocks  buffy  olive  or  buffy  brown.  A  few 
dark  spots  on  under  parts.  Throat  not  different  from  breast.  Iris  chrysolite 
green  above  and  below  with  black;  black  through  the  middle  of  the  eye.  Iris 
rim  cream-buff. 

Another  male.  Sides  and  upper  parts  dark  olive-buff.  Upper  parts  citrine 
becoming  sulphine  yellow,  olive  lake,  or  ecru-olive,  then  reed  yellow  or  prim- 


TOADS:  BUFONIDAE 


I  j  I  j  1 1    B.b.  halophilus 

III 


Map  n 

rose  yellow,  then  cartridge  buff  on  lower  side.  Some  warts  with  honey  yellow 
on  hind  legs  and  sides;  browning  olive  on  warts  along  median  line.  Median 
line  reed  yellow  from  snout  to  vent.  Under  parts  pale  olive-buff.  Throat  with 
wash  of  cream-buff.  Buttocks  and  lower  belly  Isabella  color  and  purplish. 
Black  areas  of  upper  parts  with  buckthorn  brown  centers.  A  few  spots  of 
ochraceous-orange  along  sides. 


152  HANDBOOK  OF  FROGS  AND  TOADS 

Female.  Grayish  olive  above;  upper  parts  of  fore  and  hind  legs  light  grayish 
olive;  or  olive-buff  upper  parts;  or  light  mineral  gray  on  all  upper  parts.  Stripe 
down  back  chalcedony  yellow  or  citron  yellow.  Upper  parts  with  numerous 
black  areas;  these  often  with  olive-buff  centers.  Along  either  side  of  median 
line  are  warts,  some  with  dark  olive-buff  or  picric  yellow  centers.  Throat  olive- 
buff,  pectoral  region  and  upper  belly  pale  olive-buff.  Under  part  of  femur  and 
lower  belly  olive-gray  or  deep  olive-gray.  Iris,  sulphur  yellow  pupil  rim,  above 
which  are  black  reticulations,  then  sulphur  yellow  becoming  pale  yellow- 
green,  then  beryl  green  rim  on  outside  above,  but  seafoam  yellow  or  ivory 
yellow  behind  and  in  front.  Lower  part  of  iris  black  with  a  little  cream-buff  or 
pinkish  buff. 

Structure:  No  cranial  crests  (except  occasionally  in  very  large  individuals) ; 
parotoids  elongate,  widely  separated;  two  long  metatarsal  tubercles,  inner 
with  a  free  blunt  end;  spread  of  hind  foot  usually  less  than  36  per  cent  of  head 
and  body  length;  glands  on  tibia  present;  no  external  vocal  sac  apparent  in  the 
males;  interorbital  space  only  slightly  greater  than  internasal  space;  first  and 
second  fingers  equal;  not  so  heavily  pigmented  as  B.  boreas  boreas. 

Voice:  Its  song  is  a  slow,  deep-toned,  prolonged  trill. 

"The  males,  while  in  the  water,  utter  a  series  of  low  mellow  tremulous  notes. 
In  chorus  the  notes  may  be  compared  to  the  voicings  of  a  brood  of  young  do- 
mestic goslings.  The  call  of  each  male  is  uttered  for  a  second  or  two  and  re- 
peated at  short  intervals,  so  that  a  practically  continuous  chorus  issues  from 
a  breeding  colony.  To  human  ears  the  notes  lack  carrying  power  and  can 
scarcely  be  thought  to  be  of  use  in  attracting  toads  at  any  great  distance  from 
the  pools  where  the  males  are  calling.  Occasionally  males  in  their  daytime 
retreats  will  utter  the  notes  once  or  twice.  This  species  does  not  have  an  en- 
larged vocal  pouch  such  as  is  possessed  by  many  species  of  toads  (for  example, 
americanus,  cognatus,  tvoodhousii)^  and  this  lack  of  a  'resonating  pouch*  is 
probably  responsible  for  the  small  volume  of  sound  uttered.  Calling,  with 
halophilus,  is  to  be  heard  in  the  daytime  as  well  as  at  night"  (T.  I.  Storer, 
Calif.,  1925,  p.  177). 

Breeding:  This  species  breeds  from  January  to  July,  according  to  the  climate 
of  location.  The  eggs  are  in  long  strings  laid  at  the  margins  of  ponds  or  at 
edges  of  flowing  streams  and  are  occasionally  in  two  or  three  rows.  There  are 
no  partitions  between  the  eggs.  The  vitellus  is  Mo  inch  (1.7  mm.),  the  outer 
tube  %  inch  (5.0  mm.),  the  inner  tube  }4  inch  (3.6  mm.).  The  dull  blackish 
medium  tadpoles  are  2%  inches  (55  mm.).  The  tooth  ridges  are  %.  After  28 
to  45  days,  the  tadpoles  transform  from  April  to  August  at  %-%  inch  (12-15 
mm.). 

Journal  notes:  Feb.  12, 1942.  Visited  W.  M.  Ingram.  He  had  a  pair  of  B.  bo- 
reas halophilus  mated  axillary  fashion. 

March  6.  About  10  A.M.  Anna  and  I  went  to  Stanford  artificial  lake  to  look 
for  A.  t.  calif orniense.  Didn't  find  any.  At  last  found  at  end  of  a  pond  a  mated 


TOADS:  BUFONIDAE 


'53 


Plate  XXVIU.  Bufo  boreas  halophiltts  Plate  XXIX.  Bufo  boreas  nelsoni  (X%). 

.  i.  Male.  2,3.  Females.  1,2,3,6.  Males.  4,5.  Females. 


154  HANDBOOK  OF  FROGS  AND  TOADS 

pair  of  Bufo  b.  halophtlus.  The  male  has  a  friendly  chuckle,  which  it  gives  at 
times  while  mated.  Presently  I  came  to  an  area  in  the  water  near  a  fallen  tree. 
Here,  all  about,  at  least  10  feet  out  from  the  bank  in  water  12  inches  to  2  feet 
deep,  were  long  strings  of  eggs.  The  pairs  must  be  very  restless.  The  strings 
are  wound  around  many  tussocks  and  are  often  in  two  bands  for  a  long  dis- 
tance. The  spider  web  is  quite  different  from  our  usual  Bufo  a.  amcricanus 
bands,  which  are  in  the  very  shallow  muddy  edges  and  very  heavily  massed* 
The  egg  file  is  a  double  row  of  eggs  and  somewhat  like  that  of  Bufo  joivleri 
and  Bufo  woodhottsu.  Later  near  edge  saw  a  pair  "scrunch"  down  under  the 
edge  of  a  board  and  beside  a  tussock.  Captured  them  and  another  single  male. 

April  u,  around  Chico,  Calif.  In  the  forenoon  went  out  beyond  golf  links. 
Found  toad  eggs.  Later  in  grass,  beside  a  reservoir  took  a  toad.  In  edge  of  a 
stream  saw  a  toad  "scrunched"  down  in  grass.  Caught  it.  In  pools  around 
reservoir  no  end  of  tadpoles.  Some  Bitfo  just  hatched. 

Authorities9  corner: 
H.  C.  Yarrow,  Nev.,  1878,  p.  208. 
J.  Grinnell  and  T.  I.  Storer,  Calif.,  1924,  pp.  655-656. 
J.  R.  Slevin,  B.C.,  1928,  pp.  95-96. 

Amargosa  Toad 

Bufo  boreas  nelsom  Stejneger.  Plate  XXIX;  Map  n. 

Range:  Southern  and  eastern  Nye  County  and  northern  Lincoln  County, 
Nev.,  to  Resting  Springs,  Morans,  and  Lone  Pine,  Owen's  Valley,  Calif. 

Habitat:  "So  far  known  from  three  separated  localities,  but  most  character- 
istic population  is  in  the  upper  part  of  the  Amargosa  River.  Apparently  this 
toad  is  more  closely  restricted  to  water  than  even  its  near  relatives  which  in- 
habit more  humid  districts"  (J.  M.  Linsdalc,  Nev.,  1940,  p.  204). 

Size:  Adults  i%~3%  inches.  Males  42-68  mm.  Females  46-89  mm. 

General  appearance:  "Similar  to  B.  boreas:  skin  between  warts  smooth; 
snout  protracted,  pointed  in  profile;  webs  of  hind  legs  very  large;  soles  rather 
smooth;  limbs  shorter,  elbows  and  knees  not  meeting  when  adpresscd  to  the 
sides  of  the  body;  inner  metacarpal  tubercle  usually  very  large"  (L.  Stejneger, 
Calif,  and  Nev.,  1893,  p.  220). 

Color:  Springdale,  Nev.,  May  13,  1942.  This  is  another  boreas  type  toad. 
The  general  background  of  back  is  buffy  olive,  with  the  side  of  face  and  upper 
part  of  hind  limbs  the  same.  The  stripe  down  mid-back  is  primrose  yellow  to 
pale  olive-buff.  The  parotoids  bear  the  most  prominent  color  of  the  back,  being 
tawny-olive  to  tawny.  There  are  three  paired  rows  of  tubercles:  one  along 
either  side  of  middorsum,  one  from  parotoid  backward,  and  the  third  an  ir- 
regular intermediate  one.  All  these  tubercles  have  centers  of  cinnamon-brown 
or  mikado  brown.  Back  of  the  angle  of  mouth  is  a  patch  of  ochraceous-tawny 
or  ochraceous-orange,  which  color  extends  backward  topping  a  row  of  very 


TOADS:  BUFONIDAE  155 

flat  tubercles.  Below  these  the  black  spots  become  large  and  prominents  each 
centered  with  a  light-tipped  tubercle.  There  is  a  prominent  black  area  below 
the  eye,  another  from  tympanum  downward,  and  sometimes  one  below  the 
nostril,  all  three  along  the  upper  labial  border.  The  upper  eyelid  bears  some 
tawny-olive  or  tawny.  The  iris  is  black  with  a  cream  color  or  citron  yellow 
pupil  rim,  the  same  colors  flecking  the  whole  eye.  There  is  a  prominent 
oblique  bar  of  primrose  yellow  or  marguerite  yellow  at  the  front  edge  of  eye. 
The  throat  is  primrose  yellow  to  marguerite  yellow  almost  clear  of  spots;  the 
rest  of  under  parts  arc  pale  olive  with  scattered  black  spots. 

Males  and  females  are  alike  in  color,  but  after  being  in  captivity  and  after 
breaking  their  axillary  mating  hold  the  males  were  very  black  and  the  female 
markedly  lighter,  this  past  night.  When  first  brought  in  all  five  toads  looked 
quite  light.  This  afternoon  they  are  much  the  same  in  color  pattern. 

Structure:  "The  51  specimens  now  on  hand  from  Oasis  Valley  and  the 
Amargosa  River  near  Beatty  .  .  .  contrast  with  toads  of  the  species  Bnjo  boreas 
to  the  north  and  west  in  the  following  characters:  Small  size:  the  largest  adult 
in  the  lot  measures  72.5  mm.  in  head  and  body  length.  This  is  a  little  more 
than  half  the  maximum  size  of  Bufo  b.  boreas.  The  narrow,  wedge-shaped 
head,  especially  when  viewed  from  below,  is  one  of  m the  striking  peculiarities 
of  this  toad.  This  seems  to  be  correlated  with  the  snout  protracted  and  pointed 
in  profile  as  mentioned  by  Stejneger.  Limbs  so  reduced  that  when  adpressed 
to  sides  of  body  elbows  and  knees  do  not  meet,  as  was  pointed  out  by  Stejneger. 
Small  feet  and  reduced  webbing  are  especially  noticeable  and  are  just  the  re- 
verse of  Stejneger's  statement,  which  may  not  have  been  as  he  intended.  Re- 
duced spots  below  in  all  the  large  individuals  contrasts  with  boreas  and  with 
all  small  ones  from  the  Amargosa  Valley.  Possibly  the  old  ones  tend  to  lose 
ventral  spotting.  Smooth  skin  and  small  weakly  developed  warts  which  char- 
acterize this  lot  may  be  an  indication  of  close  restriction  of  these  toads  to  the 
water.  Inner  meiacarpal  tubercle  appears  to  average  large,  as  Stejneger  indi- 
cated" (J.  M.  Linsdalc,  Ncv.,  1940,  p.  204) . 

Voice:  Beatty,  Nev.  The  male  frequently  gives  a  short  cheerful  chuckle.  We 
don't  know  that  we  can  tell  this  from  the  chuckle  of  B.  exul.  Both  are  boreas 
types. 

Breeding:  J.  M.  Linsdale,  D.  M.  Hatficld,  and  J.  K.  Doutt  secured  trans- 
forming toads  in  May  from  15-18  mm.  in  length. 

"Near  Beatty,  about  May  20, 1931,  many  small  young  toads  were  picked  up 
near  the  stream;  large  tadpoles  were  obtained  here  on  May  4,  1936"  (J.  M. 
Linsdale,  Nev.,  1940,  p.  204). 

Journal  notes:  May  13, 1942.  At  Springdale,  Nev.,  in  the  hamlet  itself,  saw  a 
muddy  pool  with  full-grown  tadpoles;  looked  around  but  saw  no  toads.  Came 
to  Beatty  about  4  P.M.,  walked  i%  miles  north  along  Amargosa  River.  No 
toads.  Went  to  Springdale.  In  her  pool,  Mrs.  Howard  said  they  had  tadpoles. 
I  looked  at  them.  They  were  about  one-third  grown.  She  said  the  eggs  were 


156  HANDBOOK  OF  FROGS  AND  TOADS 

laid  two  weeks  ago  in  the  water-lily  box.  "They  were  in  strings  with  a  little 
egg,  then  space,  then  another  egg."  One  man  said  these  toads  could  be  seen 
almost  any  night. 

Last  night  after  8  we  went  from  Beatty,  Nev.,  to  Springdale.  With  dogs 
barking  and  peacocks  calling  we  thought  our  mission  ruined.  We  went  to 
our  tadpole  pool.  Across  the  pool  I  saw  a  light  toad  or  frog.  Made  for  it  and 
caught  my  first  Bufo  b.  nelsoni,  the  Amargosa  toad.  Soon  saw  another  resting 
in  shallow  water.  Went  to  another  pond  near  by  and  here  soon  espied  two 
more — a  large  male  and  a  small  female.  None  are  mated.  These  are  shallow 
pools,  mucky,  and  no  more  than  2  to  8  inches  deep.  Returned  to  the  first  pool 
and  here  another  male  was  suspended  in  water  in  the  weeds.  These  are  not 
hard  to  capture.  Where  were  they  in  the  daytime  when  I  was  here?  Last  night 
about  5,  at  the  suggestion  of  Mrs.  Howard,  we  looked  up  Bobbie  Shoshone, 
an  alert  Indian  boy,  and  told  him  what  we  wanted. 

May  14.  Bobbie  Shoshone  went  out  last  night  and  caught  one  Bufo  b.  nel- 
soni and  several  Hyla  regtlla.  This  morning  he  went  out  again  and  secured 
three  more  toads. 

Authorities9  corner: 

L.  Stejneger,  Calif,  and  Nev.,  1893,  pp.  220-221. 
J.  M.  Linsdale,  Nev.,  1940,  p.  204. 

Southern  California  Toad,  Arroyo  Toad 

Bufo  calif ormcus  (Camp).  Plate  XXX;  Map  8. 

Range:  Coastal  region  of  southern  California — San  Luis  Obispo  Co.  (Miller 
and  Miller,  1936)  through  Ventura  Co.  (Camp,  1915)  and  San  Diego  Co. 
(Klauber,  1928) — to  lower  California  on  Rio  Santo  Domingo  (Klauber,  1955; 
Tevis,  1944). 

Habitat:  Arroyo  stream  beds;  inland  valleys  and  foothills. 

"In  San  Diego  County  the  range  seems  largely  restricted  to  the  sandy 
washes  of  the  rivers  in  the  Upper  Sonoran  Zone"  (L.  M.  Klauber,  Calif.,  1931, 
p.  141). 

"The  river  bottom  has  marginal  growths  of  oaks  and  cottonwoods  on  sandy 
beaches  with  willow  and  Bacchans  thickets  bordering  the  stream.  The  small 
flow  of  clear  water  gave  promise  of  continuing  well  through  the  summer.  The 
stream  subdivided  frequently  and  at  places  formed  comparatively  quiet  pools 
in  the  gravel  as  much  as  18  inches  deep.  It  was  within  a  few  feet  of  one  of 
these  that  a  trilling  animal  was  watched  in  the  beam  of  a  flashlight  as  it  sat 
half  submerged  in  a  puddle  not  more  than  2  feet  across.  The  fore  part  of  the 
throat  was  moderately  distended,  not  to  compare  with  the  Hyla  rcgilla  which 
were  calling  on  all  sides"  (L.  and  A.  H.  Miller,  Calif.,  1936,  p.  176). 

Size:  Adults,  i%-2%  inches.  Males,  44-58  mm.  Females,  44-68  mm. 

General  appearance:  This  is  like  a  small  Great  Plains  toad  (Bufo  cognatus) 


TOADS:  BUFONIDAE 


157 


but  is  more  uniform  in  color  on  the  back.  This  little  toad  is  very  short  and 
thick  in  body,  very  short  in  head,  and  the  arms  and  legs  appear  very  stolid. 
The  foot  is  a  little  longer  than  in  B.  cognatus.  The  eyelids  and  parotoids  are 
tuberculate.  Several  of  the  larger  warts  have  spiny  tips,  in  some  cases  several 
tips  to  one  wart.  In  a  toad  that  has  recently  shed  its  skin,  there  is  a  row  of  con- 
spicuous light  tubercles  on  the  side,  bordered  above  and  below  by  irregular 
black  lines.  In  the  young  the  light  and  dark  areas  of  the  head  and  back  are 
strongly  contrasted.  Since  1930  we  have  seen  most  of  the  Bufo  californicus 
specimens  in  collections.  Those  who  know  it  best  wonder  if  it  should  be  com- 
pared to  Bufo  compactilis  rather  than  Bufo  cognatus.  Somehow  we  cannot 
now  bring  ourselves  to  place  woodhoitsii,  w.  fowleri,  or  californicus  in  the 
lower  f  Ar00Mappet  group  of  Bufo  compactilis  and  Bufo  cognatus. 

"A  toad  with  divergent  head  crests,  a  nasal  boss,  short  slightly  divergent 
parotoids  and  with  an  internal,  cutting  tubercle  on  hind  foot;  femur  short  as 
in  Bufo  cognatus  cognatus,  the  Great  Plains  toad.  Size  medium;  parotoids 
wide;  coloration  nearly  uniform,  without  large  spots;  no  vertebral  streak;  ex- 
ternal tubercle  on  hind  foot  small  and  rounded,  not  provided  with  a  cutting 
edge"  (C.  L.  Camp,  Calif.,  1915,  p.  3?i). 

Color:  Females.  Warner's  Hot  Springs,  San  Luis  Rey  River,  Calif.,  from 
L.  M.  Klauber,  described  June  12,  1929.  Upper  parts  olive-brown  or  dark  olive. 
On  top  of  snout  are  two  parallel  black  bars  to  level  of  front  of  eye  where  they 
connect  with  transverse  bar.  From  middle  of  upper  eyelid  an  oblique  back- 
ward bar  of  black  extends  inward  but  does  not  meet  its  fellow.  Just  inside  the 
front  end  of  parotoid  is  faint  black  spot  centered  with  a  wood  brown  or  avel- 
lancous  wart.  Opposite  back  end  of  parotoid  two  narrow  black  spots  meet  at 
an  angle  pointed  forward.  Just  back  of  each  of  these  and  farther  from  meson 
is  a  larger  round  spot.  About  midway  from  eye  to  vent  is  a  pair  of  spots  near 
the  meson.  Thereafter  to  vent  spots  are  very  small,  possibly  three  or  four  either 
side  of  meson.  Ahead  of  oblique  black  bar  of  eyelid  is  a  prominent  light  band 
across  top  of  head,  olive-buff  in  young  or  wood  brown  to  avellaneous  in  adult. 
Same  color  on  front  third  of  parotoid,  between  angle  spots  and  back  of  next 
pair  of  spots  to  each  side  of  meson  midway  from  vent  to  eye.  On  each  side  an 
oblique  line  of  olive-buff  warts  extends  to  the  groin  with  black  line  below  it 
and  sometimes  an  avellaneous  band  above  with  a  black  line  above  that.  Sides 
olive-buff  or  pale  olive-buff.  Under  parts  white.  Throat  pale  olive-buff  or  til- 
leul  buff.  Underside  of  feet  and  forefeet  vinaceous  buff.  Underside  of  hind 
legs  and  rear  belly  light  vinaceous  fawn  or  light  grayish  vinaceous.  Two  or 
three  olive-buff  warts  back  of  angle  of  mouth.  Two  black  crossbars  on  fore- 
arm, none  on  brachium.  One-half  crossband  on  front  of  femur  and  two  on 
rear  half  of  tibia.  Each  bar  has  centers  or  warts  of  avellaneous  or  wood  brown. 
Space  between  tibial  crossbars  olive-buff  or  pale  olive-buff.  Dark  spot  below 
eye;  oblique  olive-buff  area  in  front  of  eye  in  young,  wood  brown  in  adults. 
Iris  black.  Pupil  above  and  below,  but  not  back  or  front,  with  prominent  rim 


158  HANDBOOK  OF  FROGS  AND  TOADS 

of  light  or  pale  chalcedony  yellow  and  with  spots  of  this  color  or  apricot  buff 
on  iris. 

Male.  Glen  Lonely,  San  Diego  Co.,  Calif.,  from  L.  M.  Klauber  and  L.  Cook, 
March  28, 1930.  The  two  males  in  hand  have  the  light  band  between  eyes,  deep 
olive-buff  or  yellowish  citrine.  The  front  third  of  parotoids  is  not  conspicu- 
ously different  from  the  rest  of  parotoid  as  in  female  described.  It  may  have 
touch  of  vinaceous  buff.  The  upper  parts  of  these  males  are  dull  citrine,  gray- 
ish olive,  or  dark  grayish  olive.  The  prominent  warts  are  tipped  with  one  to 
several  spines.  Sometimes,  but  not  always,  these  are  dark  or  brownish  tipped. 
The  dorsal  spots  of  these  males  are  not  so  prominent  as  in  females  described. 
The  short  wood  brown  or  avellaneous  band  down  mid-back  is  inconspicuous 
in  these  males.  The  lateral  oblique  line  of  olive-buff  warts  is  not  conspicuous 
in  these  males  nor  in  the  female  in  hand.  (Another  female  with  freshly  shed 
skin  is  much  brighter  in  color  and  this  line  of  light  warts  appears.)  The  throat 
is  slightly  different  from  the  rest  of  under  parts.  The  middle  and  lower  throat 
is  vinaceous-ibuff.  On  upper  jaw  below  the  rear  of  eye,  in  both  males  and 
females,  is  a  light  vertical  band  of  wood  brown,  avellaneous,  or  vinaccous-buff. 
Iris  of  each  male  is  more  spotted  and  not  so  black  as  in  the  female  in  hand. 

Structure:  Parotoids  wider  and  longer  (width  in  length  1.75-2.0)  than  in 
B.  cognatus  (width  in  length  1.5-1.8) ;  parotoid  interval  wide,  twice  the  gland 
width.  Occasionally  a  postorbital  bar  present  and  the  posterior  end  of  the  cra- 
nial valley  slightly  suggesting  embossment.  One  cutting  metatarsal  tubercle 
and  one  small  one;  femur  short;  foot  about  one-half  webbed;  hind  limbs  long; 
half  or  moie  of  femur  free  from  body  skm. 

Description  of  type:  "Size  medium;  hind  legs  very  short,  femur  almost  en- 
tirely enclosed  in  skin  of  abdomen;  head  short  and  thick;  nasal  region  elevated 
into  a  bony  protuberance;  longitudinal  cranial  crests  more  or  less  united  across 
median  region,  and  slightly  divergent;  transverse  crests  divided  by  width  of 
median  groove;  parotoids  oval,  slightly  divergent  and  very  broad;  inner  tu- 
bercle of  hind  foot  with  a  sharp  edge;  outer  tubercle  very  small,  rounded  and 
without  cutting  edge;  eyelids  and  back  evenly  tuberculated;  tympanum  oval, 
shorter  than  diameter  of  eye;  a  dozen  or  more  large  whitish  tubercles  below 
and  just  posterior  to  the  tympanum"  (C.  L.  Camp,  Calif.,  1915,  p.  332). 

Voice:  "Three  animals  were  trilling,  spaced  about  equally  along  a  quarter- 
mile  of  stream  bed.  .  .  .  There  is  considerable  resemblance  to  the  louder, 
more  raucous  call  of  cognatus,  even  granting  the  difference  which  Myers  em- 
phasizes" (L.  and  A.  H.  Miller,  Calif.,  1936,  p.  176). 

"The  call  of  cognatus  is  a  trilled  rattle,  with  much  of  the  timbre  of  Acns  in 
it.  The  call  of  calif  ornictts  is  a  sweet  trill  reminding  one  of  B.  americanus  but 
somewhat  lower  and  less  prolonged"  (G.  S.  Myers,  Calif.,  1930,  pp.  74-75). 

"The  call  is  a  beautiful,  penetrating  trill  with  a  peculiar  ventriloquistic  qual- 
ity which  makes  it  difficult  to  determine  the  direction  from  which  it  ema- 
nates" (L.  M.  Klauber,  Calif.,  1934,  p.  6). 


*59 

Breeding:  May  and  June. 

We  need  more  descriptions  of  the  egg  masses,  individual  eggs,  tadpoles,  and 
transformation. 

Journal  notes:  March  25, 1930.  Our  experiences  with  these  toads  in  life  have 
been  entirely  with  material  sent  us  by  L.  M.  Klauber.  At  present  we  have  four, 
two  males  and  two  females  in  our  toad  garden.  During  the  day  they  keep 
themselves  buried  in  the  moist  soil  with  just  their  heads  out.  They  are  not 
always  sleeping,  however,  as  they  promptly  stir  if  an  ant  is  thrown  near  them. 
Toward  evening  they  are  more  active  and  hop  around.  One  toad  seems  to 
have  adopted  an  evening  perch  in  a  pot  of  herb  robert.  During  the  evenings 
the  males  sometimes  give  brief  calls,  just  enough  to  suggest  their  little  trill. 

April  24, 1942,  Atascadero,  Calif.  Went  to  Salinas  River  northeast  of  Santa 
Margerita  for  toads,  hoping  for  Bufo  californicus.  No  luck! 

April  25.  Big  Pines  Area:  a  public  camp  just  beyond  (E.)  of  Jackson  Lake. 
Came  to  shallow  pool  beside  the  road,  shadowed  with  willows.  The  outlet  was 
through  grassy  field  and  row  of  willows  and  shortly  fell  rapidly  into  a  deep 
cut.  While  Bert  explored  the  willow  tangle  outlet,  I  walked  slowly  around 
edge  of  pond.  There  were  tussocks  of  grass.  I  soon  came  to  a  shallow  corner 
and  there  tangled  on  surfaces  of  grasses  were  two  toad  egg  strings.  These 
seemed  small  and  narrow  for  Bufo  b.  halophilus.  Eggs  close  set  and  tending 
to  be  in  a  double  row,  type  like  Bufo  woodhousii  or  Bufo  fowleri.  Are  these 
Bufo  californicus  or  Bufo  b.  halophilus?  Not  the  same  as  Bufo  b.  halophilus 
we  saw  at  Palo  Alto.  If  they  are  Bufo  californicus  eggs  they  are  totally  unlike 
Bufo  compactilis  or  Bufo  cognatus  eggs. 

May  5,  Have  we  arrived  at  Klauber's  "3%  or  4  miles  south  of  Buckman 
Springs"  where  he  caught  Bufo  calif ormcus,  i.e.,  at  the  bridge  below  the  con- 
fluence of  La  Posta  and  Cotton  wood  Creek  ?  Here  is  a  wide  expanse  of  sand, 
willow  along  creek,  oaks  farther  back.  We  took  here  three  T.  o.  hammondii, 
two  H.  regilla,  and  one  R.  a.  draytonii.  This  would  be  an  ideal  place  to  get  the 
toad  if  it  were  out.  Saw  no  Bufo,  Rana,  or  Hyla  regilla  tadpoles. 

May  6.  Collected  at  Green  Valley  Falls  Public  Camp  on  Swcetwater  River 
(creek).  Just  above  water  crossing,  found  two  fresh  complements  of  Bufo. 
Each  is  a  small  complement  of  a  small  toad.  They  are  more  or  less  in  double 
arrangement  in  tube  like  Bufo  woodhousii,  Bufo  w.  fowleri,  Bufo  b.  boreas, 
and  Bufo  b.  halophilus.  Two  other  complements  are  hatched. 

Robert  Livezey  of  Stockton,  Calif.,  has  examined  these  eggs.  He  concludes 
that  they  are  different  from  Bufo  boreas  halophilus  and  from  the  Bufo  boreas 
boreas  he  knew  at  Corvallis,  Ore.  We  both  suspect  they  are  Bufo  californicus, 
the  form  we  so  often  sought  in  vain.  Our  summary  follows:  "Files  or  strings 
with  a  continuous  gelatinous  encasing.  One  envelope  present.  Envelope  more 
than  5  mm.  in  diameter;  5.62-6.12  mm.,  average  5.77  mm.;  distinct  and  rela- 
tively firm.  Vitcllus  1.25-1.62  mm.,  average  1.42  mm.;  black  above  and  gray 
below.  Average  of  42  eggs  per  inch;  several  thousand  per  female.  Deposited 


160  HANDBOOK  OF  FROGS  AND  TOADS 

in  tangled  strings  on  bottom  of  pool  among  leaves,  sticks,  gravel,  mud,  etc. 
Range,  coastal  area  of  southern  California  from  upper  Salinas  Valley  to  Lower 
California  on  Rio  San  Domingo.  Season  May  and  June.  Bufo  californicus" 
We  hold  that  Bufo  cognatus,  B.  b.  boreas,  and  Bufo  b.  halophilus  have  two 
envelopes  in  a  file.  Our  identification  of  these  eggs  as  B.  californicus  is  pre- 
sumptive and  circumstantial. 

Authorities9  corner: 
C.  L.  Camp,  Calif.,  1915,  p.  332. 
G.  S.  Myers,  Calif.,  1930,  pp.  75-76. 
L.  Tevis,  Jr.,  L.C.,  1944,  p.  6. 

Yosemite  Toad,  Yosemite  Park  Toad 

Bufo  canortts  Camp.  Plate  XXXI;  Map  n. 

Range:  Yosemite  National  Park,  high  central  Sierra  Nevada,  7,000-11,000 
feet  altitude.  West  branch  of  McGee  Creek  trail,  Mono  Co.,  Calif.,  at  10,500 
feet,  Sept.  18, 1939  (O.  R.  Smith  and  K.  Stanton).  Two  stations  in  Tuolomne 
County,  35  miles  northwest  from  Tioga  Lake  locality  (I.  L.  Wiggins,  Calif., 
1943,  p.  197).  Occupies  the  Canadian  and  Hudsonian  life  zones,  extending 
even  into  Alpine-Arctic  (J.  Grinnell  and  C.  L.  Camp,  Calif.,  1917,  pp.  143- 
144) .  (See  Map  i,  p.  6.) 

Habitat:  Wet  mountain  meadows,  margins  of  streams  and  lakes. 

Size:  Adults,  r%-3  inches.  Males,  46-64  mm.  Females,  45-75  mm. 

General  appearance:  This  small  toad  has  a  moist  skin  and  the  skin  is  smooth 
between  the  few  large,  low-rounded  warts.  The  large  broad  parotoids  marked 
with  russet  pattern  or  crossed  by  the  pattern  of  the  back  and  the  glands  on 
legs  are  the  conspicuous  marks.  This  species  is  dimorphic,  the  female  with 
conspicuous  touches  of  black,  yellow,  and  russet,  the  male  somber  olive. 

Color:  Male.  Described  Oct.  12,  1929.  The  male  lacks  the  striking  pattern 
of  the  female,  being  an  almost  uniform  yellowish  olive  to  olivaceous  black 
with  a  broken  thread  of  yellow  down  midback.  The  black-dotted  borders  of 
the  darker  markings  and  dark  areas  around  warts  are  evident  to  greater  or 
less  degree  as  the  toad  is  lighter  or  darker  in  color.  These  may  be  outlined  by 
broken  threads  of  yellow — a  delicate  tracery.  Along  the  sides  where  belly 
merges  into  side  may  be  a  yellowish  band. 

Female.  Yosemite  Park,  Calif.,  from  C.  A.  Harwell  and  Dr.  H.  C.  Bryant. 
Beginning  with  upper  eyelid  there  extends  along  the  back  either  side  of  the 
narrow  vertebral  line  a  longitudinal  band  of  black.  In  this  there  are  from  time 
to  time  warts,  dark  olive-buff  or  isabella  color.  On  rest  of  back  are  large  spots 
of  black  with  these  colored  warts  in  them.  On  sides,  large  black  spots,  but  with 
few  or  no  warts  in  them.  Middorsal  stripe  may  be  pale  lemon  yellow  to  sul- 
phur yellow  or  primrose  yellow.  Background  color  is  lime  green,  citron  green, 
light  chalcedony  yellow,  sulphine  yellow,  or  olive-lake.  Back  of  angle  of  mouth 


TOADS:  BUFONIDAE 


161 


Plate  XXX.  Bufo  californicus  (X%).  1,2.  Plate  XXXI.  Bufo  canorus  (X%).  1,2,5. 

Males.  3,4.  Young.  5.  Female.  Males.  3,4.  Females. 


162  HANDBOOK  OF  FROGS  AND  TOADS 

is  a  xanthine  orange  wart.  Along  on  sides  and  base  of  arm  insertion  are  some 
mars  yellow  or  xanthine  orange  spots  on  ground  color.  Fore  limb  heavily 
spotted  with  black.  Hind  limb  to  tarsus  same,  also  on  rear  of  femur.  Edge  of 
upper  eyelid  and  rear  part  of  parotoid  xanthine  orange  or  less  bright.  From 
upper  eyelid  across  parotoid  is  a  longitudinal  band  of  pale  drab  gray.  The  yel- 
lows of  the  background  color  encircle  the  black  spots  and  in  alcohol  or  formal- 
dehyde make  them  appear  white  encircled.  Under  parts  white  or  pale  smoke 
gray  or  pale  olive  gray.  Few  spots  on  side  of  lower  belly,  and  a  very  few  on 
rear  of  buttocks  or  on  ventral  side.  All  in  all  this  specimen  is  pure  color  below. 
Iris  with  background  color  plus  xanthine  orange  or  mars  yellow;  this  reticu- 
lated with  black. 

Half-grown  specimen.  Oct.  12,  1929.  Unlike  large  female,  under  parts  are 
ivory  yellow  or  white  with  spotting  from  angle  of  mouth  across  pectoral  re- 
gion to  angle  of  mouth;  also  on  sides  of  belly  and  somewhat  on  belly  proper. 
Buttocks  and  lower  belly  light  mouse  gray.  Back  darker;  no  middorsal  stripe 
or  slight.  Spots  more  numerous.  Background  color  drab  or  light  drab  except 
where  surrounding  black  spots,  then  cartridge  bull  or  sulphur  yellow.  There 
is  hardly  any  impression  of  yellow  in  upper  part  as  in  full-grown  female.  Paro- 
toid with  prominent  antique  brown  or  "rusty-like"  spot.  Same  wart  back  of 
angle  of  mouth.  Little  touch  of  same  color  around  vent.  Sides  of  head  and  face 
spotted,  as  is  back.  Iris  black  with  some  sulphur  yellow,  pale  brownish  vina- 
ceous,  or  some  "rusty,"  but  more  black  than  full-grown  adult  female. 

Male.  Percgoy  Meadows,  Yosemitc  National  Park,  Calif.,  from  Carl  E.  Van 
Deman  and  G.  B.  Upton,  June,  1930.  These  males  vary  greatly  in  color  when 
kept  in  tub  of  dirt  and  when  exposed  to  sunlight.  The  back  may  be  olivaceous 
black  or  dark  olive,  becoming  deep  olive  on  the  sides.  The  wart  at  corner  of 
mouth  may  bear  a  dash  of  ochraceous-orange  or  orange-rufous  and  the  paro- 
toids  a  wash  of  Sanford's  brown.  There  is  a  reed  yellow  thread  down  the  back, 
broken  and  irregular.  Lower  surfaces  are  white  to  seafoam  yellow  with  many 
black  dashes  around  edge  of  jaw,  in  pectoral  region,  and  on  belly.  The  iris 
bordering  pupil  is  bull-yellow,  then  a  ring  crossed  with  network  of  black  lines 
gives  medal  bronze  appearance.  Along  the  upper  edge  is  a  band  of  kildare  or 
glass  green  specks. 

Held  in  hand  for  this  description,  the  color  has  grown  lighter,  the  back  is 
yellowish  olive,  the  sides  light  yellowish  olive.  The  bars  on  legs  appear  as 
dark  greenish  olive  bands  with  black  dots  around  them  forming  irregular  bor- 
ders. Some  dark  greenish  olive  areas  appear  on  back  surrounding  some  of  the 
warts.  Some  of  the  warts  have  wash  of  Sanford's  brown  or  amber  brown. 
There  is  a  dash  of  reed  yellow  in  front  of  eye.  After  photographing,  the  back 
had  become  yellowish  oil  green. 

Structure:  Short,  broad,  raised  parotoids  close  together;  muzzle  rounded; 
cranial  crests  lacking  or  slight  in  some  males;  legs  short;  gland  on  hind  leg; 
interorbital  space  narrow;  internasal  space  wide;  warts  few,  low,  flattened,  and 


TOADS:  BUFONIDAE  163 

rounded;  foot  with  two  metatarsal  tubercles;  skin  moist  like  that  of  a  Ran  a. 

This  toad  has  a  distinct  rounded  ridge  from  the  nostril  over  the  outer  edge 
of  the  eyelid.  This  gives  the  canthal  region  a  distinct  border  above  and  with 
the  steep  snout  gives  the  toad  a  square-snouted  appearance.  The  head  is  not 
so  short,  however,  as  in  Bufo  cognatus.  The  nostrils  are  widely  separated.  In 
general  shape,  the  toad  reminds  us  of  Bufo  boreas  halophilus.  Its  color  is  inter- 
mediate between  the  color  of  that  form  and  the  heavy  inky  blotches  and  con- 
trasting light  areas  of  Bufo  cognatus. 

"In  profile,  lack  of  head  crests,  small  tympanum,  and  short  legs  this  toad 
resembles  Bufo  boreas  and  its  subspecies,  but  may  be  distinguished  at  once 
from  these  forms  by  its  smaller  size,  enormous  width  of  parotoids,  slight  in- 
terval between  parotoids,  very  smooth  skin,  absence  of  a  broad  vertebral  stripe, 
and  markedly  different  color  pattern  in  both  sexes.  In  extent  of  webbing  of 
hind  foot  the  present  species  most  nearly  resembles  B.  boreas  halophilus 9  its 
near  neighbor  in  the  southern  Sierra  Nevada  and  the  San  Joaquin  Valley. 
Specimens  of  B.  boreas  boreas  from  Mono  County,  directly  to  the  east  of  the 
range  of  canorus,  have  the  large  hind  foot  characteristic  of  the  more  northern 
subspecies"  (C.  L.  Camp,  Calif.,  19163,  p.  60). 

Voice:  The  song  is  a  sustained,  rapid,  melodious  trill. 

"The  specific  name  selected,  canorus,  refers  to  the  long-sustained,  melodi- 
ous trill  uttered  by  this  toad.  This  diurnal  singing  accompanies  the  breeding 
activities,  which  take  place  as  soon  as  the  snow  melts  from  the  Sierran  mead- 
ows, June  i  to  July  15.  Many  of  the  females  captured  at  this  time  contained 
mature  eggs"  (same,  p.  62). 

"The  mating  song  of  the  Yosemite  toad  is  a  sustained  series  of  ten  to  twenty 
or  more  rapidly  utteied  notes,  constituting  a  'trill/  and  the  whole  song  is  re- 
peated at  frequent  intervals.  The  notes,  though  mellow  in  character,  carry 
well  considering  the  size  of  the  animals,  and  have  a  ventriloquial  quality 
which  makes  it  difficult  to  locate  any  one  animal  by  sound  alone.  When  a 
number  of  males  are  giving  their  songs  m  the  same  place  the  songs  overlap 
one  another  so  that  the  general  chorus  is  continuous.  There  is  some  difference 
in  the  pitch  at  which  the  several  members  of  a  group  sing,  varying  perhaps 
with  the  size  of  the  individual  toad.  .  .  .  The  notes  recall  the  courting  song 
of  the  Texas  nighthawk"  (J.  Gnnnell  and  T.  I.  Storer,  1924,  p.  659). 

"The  toad  chorus  at  different  levels  may  begin  at  least  as  early  as  May  20 
and  last  until  July  9;  and  according  to  our  experience,  singing  is  carried  on 
quite  through  the  daylight  hours  and  into  the  early  evening  at  least"  (J.  Grin- 
nell  and  T.  I.  Storer,  Calif.,  1924,  p.  660). 

Breeding:  These  toads  breed  during  late  spring  and  summer,  May  20  to 
July  16.  G.  S.  Myers  at  Peregoy  Meadows  June  20,  1928,  secured  transformed 
toads  10%  nim.  long  and  tadpoles  27  mm.  long  with  teeth  %.  |T.  I.  Storer, 
Calif.,  1925,  p.  42,  figures  teeth  %.] 

Journal  notes:  "June  5>  I930?  Peregoy  Meadows.  Saw  at  least  two  dozen  old 


164  HANDBOOK  OF  FROGS  AND  TOADS 

and  young.  Majority  were  young.  They  were  not  so  abundant  as  the  croaking 
tree  frogs,  Hyla  regilla.  Hadn't  distinguished  sexual  dimorphism  until  I 
found  a  mated  pair  in  a  very  wet  grassy  meadow,  water  three  or  four  inches 
deep  when  one  steps  in  it.  These  meadows  are  surrounded  by  snow  fields. 
Found  no  toad  eggs.  Toads  must  have  just  come  out"  (C.  E.  Van  Deman). 

July  24,  1934.  Between  Tuolomne  and  Dana  Meadows.  An  intermittent 
stream  went  through  the  meadow  leaving  many  isolated  pools.  No  standing 
pools  over  the  whole  surface,  though  the  water  comes  almost  to  the  surface 
in  some  of  the  "lat"  holes.  Our  first  sight  of  Bufo  canorus  was  of  something 
scuttling  along  the  runway  of  a  "rat"  hole  at  the  base  of  a  shrub.  We  found 
two  small  toads  here.  Near  the  pools  we  found  many  transformed  ones,  and 
several  adults  were  in  holes  or  under  overhanging  turf  at  the  edge  of  the  pool. 
We  found  one  adult  under  a  stump.  Under  a  shrub  were  two  in  water  and 
one  on  land.  Many  tadpoles  and  transforming  toads  were  in  one  of  the  rather 
shallow  pools;  many  transforming  ones  just  under  the  edge  of  tussocks  in  the 
water. 

At  Dana  Meadows.  Occasionally  in  a  pool  with  sedges  or  vegetation  or  in  a 
more  marshy  pond,  Bufo  canorus  adults  started  to  swim  out  into  the  water. 
In  a  few  pools  may  have  seen  as  many  as  20  to  25  Bufo  canorus  tadpoles.  The 
predominant  tadpoles  are  of  Hyla  regilla. 

Authorities'  corner: 
C.  L.  Camp,  Calif.,  19163,  p.  61. 
J.  Grinnell  and  T.  I.  Storer,  Calif.,  1924,  pp.  658-659. 

Great  Plains  Toad,  Say's  Toad,  Western  Toad,  Plains  Toad,  Texas  Toad, 

Western  Plains  Toad 

Bufo  cognatus  (Say).  Plate  XXXII;  Map  10. 

Range:  Imperial  Valley,  Calif.,  up  Colorado  River  to  southeast  Nevada, 
through  Salt  Lake,  Wyoming,  Montana  to  Alberta,  thence  to  extreme  western 
Minnesota  and  Iowa,  northwest  Missouri,  western  two-thirds  of  Kansas  and 
Oklahoma  southward  to  Texas  coast;  also  in  Mexico. 

Habitat:  Grazing  lands  or  agricultural  lands  of  the  Great  Plains,  along 
irrigating  ditches,  flood  plains  of  streams,  and  overflow  bottom  lands. 

Size:  Adults,  i%-4  inches.  Males,  47-95  mm.  Females,  60-99  mm- 

General  appearance:  This  is  a  large  broad-bodied  toad  with  a  general  color 
of  brown,  gray,  or  greenish.  There  is  a  light  middorsal  stripe,  and  the  back  is 
marked  with  dark  spots.  These  spots  may  be  broken  up  and  the  light  areas  or 
borders  become  more  conspicuous  so  that  the  toad  appears  obliquely  streaked 
with  light  bars  on  the  sides.  The  spotted  condition  seems  the  more  common. 
Often  there  are  four  pairs  of  bright  green  spots  down  the  back,  the  pair  at  the 
rump  leading  diagonally  to  the  groin.  There  are  green  spots  on  the  legs,  and 
an  oblique  row  of  green  spots  extending  backward  from  the  sharply  raised 


TOADS:  BUFONIDAE  165 

parotoid.  There  is  a  green  band  along  the  side.  These  green  spots  are  partially 
outlined  with  black  on  the  green  and  cream  on  the  outer  edges.  The  under 
parts  are  light,  including  the  throat.  In  these  males  the  light  throat  extends 
back  as  a  flap  over  the  thin,  dark-colored  skin  of  the  lower  throat,  which  ex- 
tends forward  when  inflated.  In  another  toad  the  green  spots  are  more  broken 
up,  and  the  green  band  on  the  side  becomes  broken  into  spots.  The  hands  and 
feet  are  light  with  dark  tips. 

Color:  Male.  Bard,  Imperial  Co.,  Calif.,  from  L.  M.  Klauber,  May  15,  1930. 
Back  drab,  buffy  olive,  buffy  brown,  light  drab,  or  avellaneous.  Parotoid  same 
color  as  back.  Fore  and  hind  legs  same  as  back,  or  lighter,  such  as  ecru-drab. 
On  sides  interstices  of  green  spots  are  pallid  brownish  drab,  pallid  vinaceous- 
drab,  or  pale  drab-gray.  Same  color  on  light  areas  on  side  of  face.  Dark  dorsal 
spots  citrine-drab,  deep  grape  green,  or  grayish  olive  outlined  by  chrysolite 
green  edges  or  deep  seafoam  green.  On  sides,  warts  of  green  spots  have  rainette 
green  or  deep  chrysolite  green  tips  and  interstices.  On  dorsum,  warts  are  wood 
brown  or  avellaneous.  Front  and  rim  of  upper  eyelid  vmaceous-fawn.  Round 
spot  on  upper  eyelid.  Elongate  pair  inside  rear  end  of  parotoid  and  near 
meson.  Another  oblique  pair  in  middle  of  back.  Anothersmall  pair  far  back 
and  near  meson,  and  another  pair  far  separated  just  ahead  of  horizontal  of 
vent  and  just  at  edge  of  femur.  From  rear  of  parotoid  a  long  spot.  From  eye 
over  tympanum  along  lower  edge  of  parotoid  a  spot.  This  broken  but  soon 
resumed  as  long  band  from  shoulder  insertion  to  groin.  Between  this  and 
mesal  spots  are  irregular  small  spots;  so  also  below  this  spot.  Fore  limbs  and 
hind  limbs  with  prominent  spots,  also  groin.  Dark  spot  from  eye  to  over 
angle  of  mouth.  Spot  below  eye.  Faint  line  from  eye  to  nostril  to  labial  edge; 
spot  oblique  below  nostril.  Under  parts  white.  Belly  and  underside  of  femur 
cream  color,  warm  buff,  or  colonial  buff,  becoming  pale  cinnamon-pink  on 
tibia.  Throat  same  color  as  venter  except  lower  throat  under  lappet,  which  is 
vivid  brown,  vmaceous-drab,  or  black.  Ins  rim  pale  cinnamon-pink  or  shell 
pink;  otherwise  black  with  much  spotting  of  light  congo  pink. 

Another  male.  Back  drab  gray  or  light  cinnamon-drab.  All  the  spots  de- 
scribed in  other  specimen  absent  or  very  small.  Face  uniform.  Toes,  fingers, 
and  membranes  somewhat  onionskin  pink  or  vinaceous-cmnamon.  Other- 
wise like  other  specimen. 

Female.  Back  buffy  brown.  Spotting  much  as  in  male.  Throat  same  color  as 
venter.  No  throat  apron. 

Structure:  Head  broad;  snout  short;  sides  and  front  of  snout  steep;  boss 
conspicuous,  ridges  divaiicating  at  rear;  parotoid  glands  small,  elevated, 
widely  separated,  oval  in  shape  extending  obliquely  downward;  two  meta- 
tarsal  tubercles  with  free  cutting  dark  edge;  hind  leg  approximates  body 
length;  hind  foot  long  in  proportion  to  hind  limb;  interorbital  space  narrow; 
internasal  space  greater  than  interorbital;  snout  only  equal  to  or  less  than  eye; 
toes  webbed,  webs  deeply  indented;  nostrils  set  far  apart;  femur  short;  horny 


166  HANDBOOK  OF  FROGS  AND  TOADS 

excrescences  on  back  of  first  finger  of  male,  and  to  less  extent  or  lacking  on 
the  second  finger. 

Voice:  The  vocal  sac  of  the  male  is  a  large  "sausage"  extending  out  and  up- 
ward. Deflated,  the  thin  discolored  skin  is  closely  folded  under  an  apronlike 
extension  of  light  colored  pebbly  skin  of  the  throat.  It  is  at  the  rear  of  the 
throat,  and  the  "apron"  may  hang  down  over  the  forebreast  as  much  as  15 
mm.  The  call  is  made  up  of  harsh,  low-pitched  notes. 

"This  form  is  one  of  the  rarer  amphibians  of  the  region.  With  Bufo  alvarms 
it  was  found  almost  always  in  the  vicinity  of  watering  troughs.  Several  were 
caught  during  the  night  of  July  7  singing  on  the  bank  of  a  pond  where  the 
large  series  of  Scaphioptis  were  taken.  .  .  .  The  song  or  trill  is  exceedingly  loud 
and  very  harsh  and  tinny,  reminding  one  very  much  of  a  Klaxon  auto  horn. 
The  interval  between  songs  is  rather  long,  nearly  a  minute,  and  as  in  Bufo 
americanus  and  Bufo  fowleri  the  trill  is  sustained  for  a  considerable  length  of 
time.  When  singing,  the  pouch  is  very  pointed  and  arises  under  the  throat 
and  projects  forward  and  upward  when  fully  distended  to  a  point  about  level 
with  the  anterior  part  of  ihe  lower  jaw  (A.  I.  and  R.  D.  Ortenburger,  Ariz., 
1926,  p.  103). 

Green  River,  Emery  Co.,  Utah.  "Songs  of  this  species  and  B.  woodhousii 
with  which  it  was  associated,  could  be  heard  in  many  parts  of  this  little  valley 
at  night"  (V.  M.  Tanner,  Utah,  1928,  p.  24). 

Breeding:  These  toads  breed  from  April  to  September,  dependent  upon 
rainfall;  in  the  northern  states  of  their  range,  from  May  to  July.  Little  is 
known  about  the  eggs  and  tadpoles.  Amplexation  pectoral;  egg  strings  single, 
sometimes  double,  no  inner  tubes  or  partitions  seemingly  of  Bufo  compactilis 
type;  mature  tadpole  mottled  gray  and  brown  dorsally;  light  greenish  yellow 
and  reddish  ventrally;  tail  fin  highly  arched,  obtuse  apex;  tooth  rows  %; 
transformation  size,  21-28  mm.;  tadpole  period  i%-2%  months  (largely  after 
A.  N.Bragg). 

Bragg's  studies  (1936,  1937?  *93Qa)  of  Bufo  cognatns  were  much  needed. 
They  mark  one  of  the  most  significant  contributions  to  anuran  life  histories, 
in  which  the  Oklahoma  group  have  recently  been  leading.  Bragg  gives  the 
individual  egg  diameter  as  1.18  mm.,  the  egg  inner  envelope  0.22  mm.  beyond 
the  egg,  and  an  outer  tube  of  jelly  2.05  mm.  in  diameter  or  2.66  mm.  when 
eggs  are  in  a  double  row.  He  gives  an  egg  complement  of  20,000,  the  eggs  with 
black  animal  pole  and  white  vegetative  pole.  The  scalloped  outer  tube  places 
this  egg  file  in  the  Bufo  compactilis  class. 

"At  hand  are  two  series  of  tadpoles  of  this  species;  one  series  was  collected 
July  2, 1938, 1.5  miles  cast  of  Meade  County  State  Park,  Kansas,  and  the  other 
lacks  data.  The  second  lot  contains  numerous  sizes  of  tadpoles  from  14  mm. 
to  31  mm.,  and  several  transforming  specimens  which  clearly  possess  the 
pattern  so  typical  of  this  species. 

"Mouthparts  in  both  series  (consisting  all  told  of  about  200  specimens)  are 


TOADS:  BUFONIDAE  167 

fairly  constant  except  in  the  transforming  and  extremely  young  specimens. 
.  .  .  The  outer  row  of  teeth  of  the  lower  labium  is  sometimes  a  little  shorter 
or  longer  than  the  figure  shows,  but  the  average  is  about  as  indicated.  The 
extent  of  the  medial  edge  of  the  papillae  on  the  lower  labium  varies  some- 
what; in  some,  the  papillae  barely  reach  the  level  of  the  ends  of  the  outer  row 
of  teeth,  while  in  others  they  overlap  the  ends  slightly. 

"Measurements  agree  with  those  given  by  Bragg,  except  that  appearance  of 
the  hind  legs  occurs  at  about  15  mm.;  the  fore  legs  appear  at  about  28  mm. 
A  pattern  recognizably  similar  to  that  of  the  adult  is  evident  at  about  20  mm. 

"These  tadpoles  show  such  a  striking  similarity  to  those  referred  by  Wright 
to  Bufo  compactilts  Wiegmann  .  .  .  that  their  conspecificity  is  suggested" 
(H.  M.  Smith,  Kans.,  1946,  pp.  Q5-96). 

Journal  notes:  July  8, 1917.  We  found  a  large  stream  rushing  through  Sierra 
Blanca,  Tex.  They  had  not  had  rain  for  six  months.  The  flat  land  was  over- 
flowed, and  a  swift  current  went  under  the  small  bridge.  At  seven  o'clock, 
while  it  was  still  day,  we  heard  no  notes  in  the  creek,  but  later  when  we  were 
camped  one-half  mile  away,  we  heard  the  chorus  plainly  and  decided  it  must 
be  spadcfoots.  .  .  .  We  found  three  species  of  toads  and  two  of  spadefoots 
migrating  from  the  mountain  sides  of  Sierra  Blanca  downward  toward  the 
pool  and  noise.  The  boys,  at  night,  captured  B.  woodhousn,  B.  cognatus,  and 
B.  compactilis  on  the  hillside  and  in  the  stream. 

May  20, 1942,  Beaver  Dam  Lodge,  Ariz.  Big  pool  north  end  of  bridge.  From 
the  pool  came  two  very  piercing  shrill  calls.  It  was  B.  cognatus.  We  collected 
the  one  we  heard  call  and  another  in  the  pool  In  the  closed  bag  its  note  is  a 
harsh  nasal  clack;  that  of  the  tvoodhottsn  is  a  friendly  chuckle. 

July  21,  El  Paso.  On  road  from  North  Junction  toward  Smeltertown  took 
one  or  two  B.  cognatus  adults  and  some  three  or  four  young. 

Authorities9  corner: 

F.  A.  Hartman,  Kans.,  1907,  p.  228.  A.  N.  Bragg,  Okla.,  19373,  p.  283. 

J.  K.  Strecker,  Jr.,  Tex.,  1910,  pp.  19-21.  G.  A.  Moore  and  C.  C.  Rigney,  Okla., 
H.  J.  Pack,  Utah,  1922,  p.  8.  1942,  p.  78. 

F.  W.  King,  Ariz.,  1932,  pp.  175-176. 

Spade-footed  Toad,  Spadefoot  Toad,  Sonoran  Toad,  Western  Toad 

Bufo  compactilis  Wiegmann.  Plate  XXXIII;  Map  8. 

Range:  Southern  portions  of  Utah  and  Nevada,  south  far  into  Mexico,  and 
east  to  southwest  Oklahoma  and  the  eastern  timber  belt  of  Texas. 

Habitat:  We  found  this  road  breeding  in  rain  pools  in  open  fields  near 
streams,  in  pools  in  creek  valleys,  in  irrigation  tanks  or  cattle  tanks.  It  is  a 
desert  form  that  may  at  times  be  seen  feeding  at  night  under  the  street  lights 
of  desert  towns. 

Size:  Adults,  2% 2-3%  inches.  Males,  52-78  mm.  Females,  54-91  mm. 


i68 


HANDBOOK  OF  FROGS  AND  TOADS 


Plate  XXXII.  Bufo  cognatus  (X%)-  M-  «<*  XXXIIL  B"/°  comfaeau.  1,2,5. 

Females.  2,?.  Males.  Males  (X%)-  3-  Male  croaking  (Xw).  4- 

Egg  files 


TOADS:  BUFONIDAE  j69 

General  appearance:  This  broad,  squatty  toad  of  medium  size  is  pinkish 
drab  in  color,  marked  with  dull  citrine  spots.  The  fingers  and  toes  are  light  in 
color.  The  under  parts  are  light.  The  back  is  covered  with  light-tipped  tu- 
bercles. 

Color:  Beeville,  Tex.,  March  27, 1925.  Male.  In  general  color  a  light  drab  or 
cinnamon-drab.  Tips  of  the  tubercles  (under  lens)  are  xanthine  orange  but 
to  naked  eye  flesh  pink,  this  giving  some  pinkish  tone  to  the  upper  parts. 
Spots  on  back,  sides,  legs,  look  dull  citrine.  Under  lens  the  spots  are  Prout's 
brown  or  some  other  brown  with  bases  of  the  tubercles  mignonette  green  and 
tips  of  tubercles  xanthine  orange.  Fingers,  toes,  and  tubercles  of  fore  and  hind 
feet  pinkish  cinnamon.  Tip  of  chin  is  white,  then  folded  part  of  throat  is 
pinkish  buff  with  ecru-olive  in  center.  This  area  is  followed  by  a  circular  pec- 
toral area  of  purplish  lilac.  Pupil  of  eye  distinctly  horizontal,  with  horizontal 
black  bar  through  the  eye  at  either  end  of  the  pupil.  Rim  cartridge  buff  or 
marguerite  yellow.  Iris  spotted  vinaceous-buff  and  bright  green-yellow  with 
sudan  brown  vermiculations.  Lower  eyelid  clearly  transparent.  Tympanum 
dark  gray. 

Female.  White  below.  Chin  with  a  few  light  grayish  olive  spots.  Tubercles, 
fingers,  and  toes  buff-yellow.  Back  is  cinnamon-drab.  Upper  parts  not  strik- 
ingly different  from  male.  Tympanum  pale  olive-gray. 

Structure:  Parotoid,  elongate,  sometimes  smooth;  no  sharp-edged  ridge 
from  eye  to  nostril,  nostril  area  smooth;  crown  without  bony  ridges,  snout 
short,  blunt;  mterorbital  space  about  equal  to  upper  eyelid;  first  finger  at  least 
equal  to  second;  toes  half  webbed;  sole  tubercles  large,  each  with  a  cutting 
edge;  tympanum  much  smaller  than  the  eye. 

Voice:  The  vocal  sac  is  a  large  fat  oblong  "sausage."  Deflated,  it  forms  a 
light  apron  covering  several  darker  folds  in  the  rear  of  the  throat.  J.  K. 
Strecker,  Jr.  (Tex.,  iQ26f,  p.  12),  calls  it  "a  loud  explosive  quack." 

May  TI,  1925.  San  Antonio,  Tex.  At  Leon  Valley  Creek  a  big  chorus  of 
B.  valliceps,  B.  debihs,  B.  compactihs,  and  S.  couchh  in  considerable  num- 
bers. .  .  .  B.  compactilis  males  were  calling  from  the  bank.  Each  one  seemed 
to  have  a  favorite  perch,  and  when  scared  away  from  it  by  our  light  some- 
times would  return  almost  immediately.  Their  call  is  loud  and  shrill  and  with 
a  big  chorus  would  be  deafening. 

May  12.  No  B.  compactilis  calling  at  Leon  Valley  Creek  tonight. 

May  29,  Comfort,  Tex.  A  few  were  calling  when  we  first  reached  the  spade- 
foot  pond.  Their  call  is  very  shrill.  ...  In  the  pond  were  many  oats  half- 
grown  or  more  with  oats  on  the  stems.  Beside  an  oat  stem  where  a  clump  of 
oats  might  be,  a  male  would  rest  in  water  in  vertical  fashion,  sausagelike 
throat  extended  i%-2  in.,  directed  outward  and  upward  above  his  head  much 
like  B.  querckus.  When  a  pair  is  picked  up,  the  male  gives  a  call  like  a  pea 
fowl  or  hen,  clucking  much  of  the  time.  The  water  was  very  shallow  where 
I  caught  the  pair. 


170  HANDBOOK  OF  FROGS  AND  TOADS 

June  15.  Left  Rio  Grande  City  7:45  A.M.  Stopped  at  Santa  Cruz  ranch  op- 
posite hill  with  U.S.  Geological  Survey  marker.  At  top  of  hill  we  could  clearly 
hear  B.  compactilis  in  a  pond  across  the  road.  These  males  which  are  croaking 
in  the  daytime  are  in  the  dense  shade  of  some  overhanging  trees. 

Breeding:  These  toads  breed  from  May  i  to  July  10,  with  a  few  stragglers 
later  at  the  time  of  the  late  summer  rains.  The  brown  and  yellow  eggs  are  in 
long  fine  coils;  the  jelly  tube  narrow,  %2  inch  (2  mm.);  the  eggs  crowded, 
14-20  eggs  in  i%  inches  (30  mm.);  and  the  vitelli  Ko  inch  (1.4  mm.).  The 
eggs  hatch  in  two  days.  The  bicolored  tadpole  is  small,  1-1%  inches  (24-28 
mm.),  light-colored,  its  back  a  drab  or  light  grayish  olive  and  its  belly  pale 
cinnamon  pink;  its  tail  crests  are  translucent.  The  tooth  ridges  are  %.  After 
40  to  60  days,  the  tadpoles  transform,  June  i  to  August  i,  at  %  inch  (12  mm.). 

Journal  notes:  March  22,  1925.  Started  for  Corpus  Christi,  Tex.,  at  6  P.M. 
Captured  B.  compactilis,  north  of  Kansas  City.  A  female  crossing  the  road. 

March  25.  Tonight  took  two  Bufo  compactilis  males.  In  no  place  are  they 
breeding  so  far  as  we  can  see.  Tonight  we  picked  up  what  we  at  first  took  for 
a  toad  of  B.  compactilis  build.  Not  until  later  did  we  realize  we  had  a  spade- 
foot. 

March  26.  Tonight  in  Beeville,  Tex.,  under  electric  light  saw  a  female  mov- 
ing along.  Caught  her. 

May  29.  Comfort  had  3  inches  of  rain  in  the  afternoon  of  May  28.  We  went 
to  San  Antonio  and  at  5:30  P.M.  started  for  Comfort.  At  Big  Joshua  Creek 
evidences  of  flood  but  no  frogs  calling.  .  .  .  Left  Boerne  and  arrived  at  Com- 
fort about  8:30  P.M.  The  Guadalupe  River  had  been  up  high  but  was  down 
at  this  time.  No  frogs  at  all.  When  we  pulled  up  to  Comfort  filling  station, 
heard  a  great  chorus  south  of  the  filling  station.  We  went  down  from  the  main 
road  south  toward  baseball  field,  then  left  toward  the  creek.  It  was  slippery. 
We  crossed  creek  and  in  a  cultivated  field  ^-%  mile  farther  on  was  a  great 
volume  of  "wows  or  meows1'  or  catcalling  (S.  cottchit).  In  this  pond  were 
many  B.  compacttlis.  A  few  were  calling  near  the  edge  but  more  in  the  water. 
...  At  the  far  end  of  the  cultivated  field  the  pond  extended  into  a  grassy 
pasture.  Here  the  B.  compactilis  were  calling  in  considerable  numbers.  A  few 
had  standings  on  the  shore,  several  were  out  in  clumps  of  grass.  They  are  well 
called  the  spadefoot  toad.  They  are  usually  the  associate  of  the  spadefoot  (S. 
couchit) . 

We  left  this  pool  and  went  on  to  Cypress  Creek  where  there  are  water  lilies. 
Nothing  doing.  We  went  on  2  or  3  miles,  then  returned.  Down  a  side  road 
heard  S.  couchii.  Found  pools  beside  the  road.  Herein  were  many  S.  couchti 
and  many  B.  compactilis. 

June  7.  As  we  approached  Encinal,  close  to  the  road  in  two  pools  were  many 
small  tads— toad  tads,  but  not  spadefoots.  We  had  lost  our  net  but  found 
strainers  good  for  catching  them.  Are  the  toads  B.  compactilis  or  B.  debtlis? 
When  the  pools  begin  to  dry  up,  gracklcs  go  there  to  eat  up  tads. 


TOADS:  BUFONIDAE 


171 


June  9.  At  Dolores,  Tex.,  collected  many  Couch's  spadefoots  and  B.  com- 
pactilis  in  large  concrete-bottomed  irrigation  tank.  Around  the  edges  are  a 
few  places  where  mud  has  gathered,  and  in  such  places  B.  compacttlis  has  dug 
into  the  mud — sometimes  leaving  an  opening,  sometimes  just  leaving  the 
earth  rough. 

June  2,  1942.  Beyond  Rockville  in  Virgin  River  saw  countless  small  Bufo 
tadpoles.  Two  to  three  miles  from  Springdale,  Utah,  I  went  after  toads  in  the 
dark.  Had  to  jump  a  drainage  ditch.  In  a  cultivated  field  no  end  of  water;  one 
pool.  I  thought  I  had  Scaphiopus  but  the  call  was  not  so  harsh — but  much 
harsher  than  the  few  Bufo  woodhousii  then  calling.  Caught  five  female 
B.  compacttlis.  When  I  approached,  the  males  stopped;  never  saw  one.  We 
saw  a  Bufo  compactihs  crossing  the  road  in  Rockville. 

July  22,  1942.  Toyahvale,  Tex.  Night  trip  west  on  highway.  Heard  one 
Bufo.  This  proved  to  be  Bufo  compactihs. 

Authorities'  corner: 

J.  K.  Strecker,  Jr.,  Tex.,  igoSb,  p.  82.  J.  K.  Strecker,  Jr.,  Tex.,  1926^  p.  12. 

J.  K.  Strecker,  Jr.,  Tex.,  1915,  p.  52.  J.  K.  Strecker,  Jr.,  Tex.,  1928^  p.  7. 

G.  P.  Englehardt,  Utah,  1918,  pp.  78-     H.  M.  Smith,  Gen.,  1947,  pp.  7-13. 

79- 

In  1940  Linsdale  (Nev.)  wrote:  "It  is  necessary,  in  making  final  taxonomic 
appraisal  of  toads,  to  study  the  living  animals  in  their  natural  surroundings. 
Until  further  field  studies  can  be  made  on  this  group  of  forms,  satisfactory 
conclusions  cannot  be  reached.  However,  the  preserved  material  is  sufficient 
to  show  that  some  changes  in  systematic  treatment  of  the  group  will  be  re- 
quired. Present  indications  are  that  these  toads  are  all  in  the  same  species.  The 
oldest  name  in  the  lot  is  compactihs.  At  least  four  races  seem  to  deserve  recog- 
nition, as  follows:  Bufo  compactihs  compactilis  Wiegmann  1833,  Bufo  com- 
pacttlis woodhousii  Girard  1856,  Bufo  compacttlis  fowlen  Hmckley  1882, 
Bufo  compactihs  cahfornicus  Camp  1915." 

He  may  not  be  far  from  the  right  solution  but  we  cannot  yet  accept  this  ap- 
praisal without  more  study  of  the  western  forms  in  the  field.  Should  Bufo 
cahfornicus  be  placed  as  a  Bufo  compactihs  subspecies  rather  than  as  a  Bufo 
cognatus  affiliate?  Is  Linsdale's  Bufo  compactihs  of  Nevada  the  same  as  the 
large  Bufo  compactihs  of  Texas?  In  1942  Dr.  Klauber  remarked  that  he 
wished  we  could  solve  the  B.  compactihs  problem.  The  Texas  Bufo  com- 
pactilis is  an  old  acquaintance,  but  the  western  representatives  we  do  not 
know  well  in  the  field. 

In  a  recent  paper  H.  H.  Smith  (Gen.,  1947)  divides  B.  compacttlis  into  two 
forms  restricting  B.  c.  compactihs  to  the  main  Mexican  plateau  and  B.  c.  spe- 
ciosus  for  the  U.S.  form.  We  are  glad  he  did  not  set  off  the  Nevada  and 
Arizona  form  as  new.  The  discontinuous  range  of  his  B.  c.  speciosus  may  not 
be  real.  Field  work  in  the  interval  in  Arizona  may  yet  produce  this  burrower. 


172 


HANDBOOK  OF  FROGS  AND  TOADS 


Little  Green  Toad,  Sonoran  Toad,  Green  Toad,  Sonora  Toad 

Bufo  debtlis  Girard.  Plate  XXXIV;  Map  12. 

Range:  "Eastern  Texas  and  Tamaulipas"  (Taylor,  1936).  "In  the  lower 
part  of  the  valley  of  the  Rio  Bravo  (Rio  Grande  del  Norte)  and  in  the  prov- 
ince of  Tamauhpas"  (Girard,  1854;  Stejneger  and  Barbour,  1938). 

Habitat:  On  grassy  mesquite  flats,  breeding  in  temporary  rain  pools, 
ditches,  or  shallow  pools  in  streams  of  intermittent  flow. 

Size:  Adults,  i-i4/f,  inches.  Males,  26-41  mm.  Females,  31.5-46  mm. 


Map  12 

General  appearance:  This  beautiful  little  green  toad  has  black  spots  on  the 
back  and  some  gold-  or  yellow-tipped  tubercles  on  the  sides  and  legs.  The 
legs  have  a  few  dark  barhke  spots  with  light  central  areas.  The  legs  are  short, 
the  snout  pointed,  the  parotoids  large  and  elongate  or  triangular,  the  body 
narrow  and  rounded  in  contour,  the  head  narrow  with  interorbital  space  pro-4 
portionally  wide.  There  is  a  black  bar  at  the  arm  insertion  and  there  may  be 
a  black  line  or  two  or  three  spots  in  the  middle  of  the  lower  breast. 

Color:  Male.  Alice,  Tex.,  March  31,  1925.  Back  spinach  or  cedar  green  or 
hellebore  green  to  calliste  green  or  cosse  green.  Back  with  black  spots  and 
black  tubercles  on  these  spots.  On  the  sides,  besides  green  and  black,  some 
tubercles  are  light  cadmium,  lemon  chrome,  or  maize  yellow.  These  colors 
also  appear  on  dorsum  of  hind  legs  and  brachium  of  fore  limb,  and  on  web- 
bing of  hind  feet  and  tubercles.  The  rear  of  thigh  is  green,  black,  and  light 
cadmium  color.  There  is  a  line  of  light  cadmium  where  dark  colors  meet  the 


TOADS:  BUFONIDAE 


'73 


white  under  parts.  Some  of  the  tubercles  on  greenish  background  with  light 
cadmium  tips.  Light  buff  or  cream-buff  along  upper  jaw  to  arm  insertion. 
Rim  of  lower  jaw  has  a  white  margin  followed  by  dense  black,  then  pale  olive- 
gray  to  deep  olive-gray.  There  is  a  black  bar  on  the  venter  in  the  pectoral 
region  opposite  arm  insertion;  also  black  on  the  brachial  insertion.  Sometimes 
there  is  a  black  line  in  the  middle  of  lower  breast  or  two  or  three  black  spots. 
Iris  is  mainly  black  with  some  pale  ochraceous-buff. 

Female.  June  15,  1925.  In  general  the  same  as  male,  but  without  the  black 
chin.  Under  parts  are  olive-buff  or  white.  In  some  the  two  pectoral  black 
spots  are  absent  or  not  so  well  developed  as  in  the  male. 

Structure:  Crown  without  bony  ridges;  snout  protruding;  first  finger 
shorter  than  the  second;  hind  limbs  usually  shorter  than  body  length;  tym- 
panum small;  toes  slightly  webbed  at  base;  foot  short;  no  tarsal  fold;  paro- 
toids  sometimes  extending  downward  to  the  level  of  the  jaw. 

In  view  of  Taylor's  (1936)  concept,  the  following  Strecker  quotation  is 
composite,  i.e.,  for  both  species:  "Structure:  Girard's  original  description 
(May,  1854)  of  Bttfo  debilts  (p.  87)  varies  so  slightly  from  his  Bitfo  insidior 
(p.  88)  that  we  include  it  for  future  clarification  of  the  status  of  each  form. 
The  description  is:  Upper  surface  of  head  without  any  crest  on  ridge.  Snout 
rounded.  Mouth  moderate.  Upper  jaw  emarginated.  Tongue  small.  Tympa- 
num small.  Parotoids  moderate  and  elongated.  Limbs  of  moderate  develop- 
ment, femur  shorter  than  tibia.  First  finger  longer  than  the  second.  A  large 
metacarpal  disk.  Toes  slightly  united  at  the  base  by  a  web.  Two  metatarsal 
processes.  No  membranous  fold  at  the  inner  lower  margin  of  the  tarsus.  Skin 
above  pustulous;  pustules  of  moderate  development;  warts  beneath  very 
small.  Color  above  brownish  yellow,  spotted.  No  dorsal  lighter  vitta.  Beneath 
of  a  uniform  soiled  yellow.  Allied  to  Bujo  speciosits.  .  .  ." 

Voice:  The  vocal  pouch  is  a  round  ball  reaching  to  the  tip  of  the  chin.  The 
call  is  cricketlike,  a  low  sustained  trill. 

March  28,  1925.  Between  San  Diego  and  Alice,  Tex.,  we  heard  a  note  abso- 
lutely new  to  us — something  like  a  cricket.  When  we  came  to  the  pond  and 
it  appeared  light-colored  under  the  flashlight,  I  knew  it  must  be  Btifo  debilis. 
It  is  rightly  named  "the  little  green  toad."  In  some  ways  its  note  makes  me 
think  of  Fowler's  toad,  but  it  is  not  nearly  so  loud  or  strong.  The  call  is  sus- 
tained. When  I  first  went  down  to  the  pond,  I  thought  I  heard  only  one,  but 
before  I  got  the  light  on  it  to  see  how  it  croaked,  two  of  them  started  hopping 
toward  the  ditch.  In  some  ways,  the  toad  and  its  note  remind  me  of  B.  qtierci- 
cusf  but  B.  quercicus  is  much  louder  and  shriller. 

When  held  in  hand  the  male  gave  a  little  batlike  click,  like  marbles  hitting 
together.  Over  in  the  mesquite,  "devil's  elbow,"  and  prickly  pear,  we  went 
for  two  or  three  we  heard,  and  came  back  with  one  of  them.  There  was  a  small 
ditch  or  runway  there. 

In  a  shallow  overflowed  mesquite  area,  several  B.  debilis  males  were  calling. 


174  HANDBOOK  OF  FROGS  AND  TOADS 

Oftentimes  when  you  approach,  or  particularly  when  you  put  light  on  them, 
they  duck  flat  to  the  ground.  Bufo  debilis  note  at  times  something  like  the 
"clucking"  one  gives  to  make  a  horse  travel  faster. 

Breeding:  This  species  breeds  from  the  last  of  March  to  mid-June.  "The 
eggs  are  in  small  strings  and  are  attached  to  grass  and  weed  stems.  .  .  .  The 
tadpoles  are  slightly  smaller  than  those  of  B.  punctatus"  (J.  K.  Strecker,  Jr., 
Tex.,  1926^  p.  10).  They  transform  at  %-%  inch  (8-11  mm.). 

Couch  at  Matamoras,  Mexico,  took  six  specimens  (USNM  no.  2621)  which 
measured  respectively  8.0,  8.5,  9.0,  10.5,  ii.o,  n.o  mm.  This  probably  repre- 
sents a  close  approach  to  transformation.  Doubtless  this  species  transforms  at 
a  little  larger  size  than  B.  quercictis,  of  which  the  smallest  transformed  indi- 
vidual recorded  is  7  mm. 

"Their  metamorphosis  is  accomplished  within  a  very  short  space  of  time. 
This  is  necessary  on  account  of  the  extremely  temporary  character  of  their 
breeding  places.  I  returned  twenty  days  later  to  one  pond  in  which  I  had 
found  debilis  breeding  and  discovered  that  it  was  almost  perfectly  dry,  only 
a  few  mud  holes  remaining  to  indicate  moisture.  In  one  of  these  mud  holes 
were  a  few  belated  tadpoles,  and  in  the  grass  along  the  banks  I  found  two 
small  toads  with  tails"  (J.  K.  Strecker,  Jr.,  Tex.,  1926^  pp.  10-11). 

Journal  notes:  April  3,  1925,  Gardendale,  Tex.  In  a  shallow  pool  near  the 
station  were  some  small  Bufo  tadpoles.  Returned  tonight  at  8  130  P.M.  Air  at 
least  72°.  Heard  3  or  4  Bufo  debtlis. 

May  9,  San  Antonio,  Tex.  At  Leon  Creek,  several  Microhyla,  and  one  Bufo 
debilis.  This  one  we  photographed.  He  did  not  duck  under  the  flashlight.  In 
the  next  valley  bottom— in  gravel  pit  stream— several  B.  valliceps,  one  Micro- 
hyla,  and  one  B.  debilis. 

May  u,  San  Antonio,  Tex.  Went  to  Somerset  in  the  afternoon  and  "spot- 
ted" ponds  by  the  roadside.  Back  to  the  city  for  supper;  then  out  Somerset 
road  again.  First  stop  on  return  7:45  at  roadside  pond  now  swelled  by  rain. 
Heard  lots  of  Pseudacns,  meadow  frogs,  Hyla  versicolor,  Bufo  valliceps.  Bufo 
valliceps  are  calling  from  temporary  islands  and  the  banks.  Little  nearer  the 
city,  beside  road  nothing  but  Microhyla.  Next  stop— roadside  pond— 25  male 
Bufo  debilis ',  sometimes  ^  or  4  together  at  edge  of  plowed  ground— 3  or  4  Man- 
cy's  garters  evidently  enemies  of  all  of  them.  Great  chorus  of  Microhyla  and 
Pseudacris. 

Jan.  18, 1942.  Visited  Al  Kirn  of  Somerset.  Showed  me  his  material— plenty 
of  Bufo  debilis.  These  he  loaned  me. 

Authorities'  corner: 

J.  K.  Strecker,  Jr.,  Tex.,  ipoSb,  p.  81.        E.  H.  Taylor,  Kans.,  1929,  p.  445. 
J.  K.  Strecker,  Jr.,  Tex.,  1915,  p.  51.  E.  H.  Taylor,  Gen.,  19363,  p.  513. 

R.  Kellogg,  Gen.,  1932,  p.  50. 


TOADS:  BUFONIDAE 


175 


Plate  XXXIV.  Bufo  debilis.  i.  Female 
(XO.  2.  Male  croaking  (X%)-  3-  M*'e 
(X%).  4,5-  Females  (X%)-  6.  Female 


Plate   XXXV.   Bufo   exsul    (X%). 
Young.  2,7,8.  Females.  3,4,5,6.  Males. 


176  HANDBOOK  OF  FROGS  AND  TOADS 

Black  Toad 
Bufo  exsul  Myers.  Plate  XXXV;  Map  n. 

Range  and  Habitat:  Known  only  from  Deep  Springs.  "Deep  Springs  Val- 
ley is  an  isolated  depression  in  the  desert  mountains  of  northeastern  Inyo 
County,  California.  It  is  elongate  in  form,  trending  northeast  by  southwest, 
about  twelve  miles  long  and  five  miles  broad  at  its  widest  part.  The  lowest 
part  of  the  valley,  at  its  wide  southwestern  end,  is  a  flat  area  of  about  three  by 
five  miles,  of  almost  exactly  5,000  feet  elevation,  although  the  rest  of  the  gently 
rising  valley  floor  is  also  very  level.  Surrounding  Deep  Springs,  the  White  and 
Inyo  Mountains  rise  to  heights  of  7,000  to  8,000  feet.  Westgard  Pass,  through 
which  one  enters  the  valley  from  Owens  Valley  to  the  west,  reaches  7,276  feet, 
and  the  top  of  the  pass  into  the  southern  arm  of  Fish  Lake  Valley,  on  the  east, 
is*  at  6,374  ^eet-  The  lowest  entrance  to  the  valley  appears  to  be  the  dry,  narrow, 
and  now  virtually  unused  Soldier  Pass,  from  the  dry  northeastern  corner  of 
Deep  Springs  Valley  into  Eureka  Valley  on  the  southeast;  the  top  of  this  pass 
appears  to  be  at  approximately  5,400  feet. 

"Like  other  desert  valleys  to  the  east  of  the  Sierra  Nevada,  Deep  Springs 
is  exceedingly  dry,  and  on  its  floor  the  vegetation  consists  of  sparse  low  desert 
brush  (Chrysothamnns}.  The  surrounding  mountains  support  growths  of 
juniper  and  pinon. 

"The  valley  has  few  sources  of  water.  Aside  from  washes  carrying  water 
only  during  infrequent  desert  rains,  I  know  of  only  three.  Wyman  Creek,  the 
course  of  which  leads  into  the  northern  end  of  the  valley,  contains  a  little  wa- 
ter, at  least  in  its  upper  reaches,  most  of  the  year,  and  tiny  Antelope  Spring, 
on  the  west  side  of  the  valley,  appears  to  be  permanent,  but  neither  of  these 
contributes  water  to  the  valley  floor,  except  during  exceptionally  heavy  rains. 
The  chief  water  source  is  formed  by  the  Buckhorn  or  Deep  Springs  which 
flow  from  the  base  of  the  southeastern  valley  wall  just  above  the  sink.  These 
springs  issue  from  the  rocks  for  a  distance  of  a  mile  or  more,  but  only  a  few 
of  them  have  a  strong  flow.  The  flow  from  the  more  southerly  springs  forms  a 
marshy  area  of  several  acres,  the  water  finally  draining  down  into  a  shallow 
lake  of  alkaline,  sulphurous  water  that  sometimes  reaches  a  diameter  of  a  mik 
or  more.  There  is  also  a  smaller  pond  of  good  water  between  the  springs  and 
the  lake.  The  marshy  area  and  the  watercourses  through  it  emit  a  strong  sul- 
phurous odor.  Much  of  this  is  apparently  due  to  sulphur  bacteria,  but  some  of 
the  springs  themselves  must  carry  sulphur"  (G.  S.  Myers,  Calif.,  1942,  pp. 

1-2). 

Size:  Adults  i%-2%  inches.  Males  44-59  mm.,  average  52  mm.,  mean  51 
mm.;  females  46-61.5  mm.,  average  54  mm.  (Measurements  of  51  specimens 
of  the  Ferris- Wiggans-Myers  accession  May  i,  1937,  and  of  the  C.  L.  Hubbs 
family  accession  Sept.  4,  1934.) 

Of  25  adults  in  our  collection  from  Deep  Springs  Valley  School,  May  12, 


TOADS:  BUFONIDAE 


177 


1942,  we  have  the  following  sizes:  adults  45-56  mm.;  males  (17)  45-53.5  mm., 
average  49  mm.,  mean  48  mm.;  females  (8)  45(?)~56  mm.,  average  49  mm., 
mean  51  mm.  There  were  five  questionable  as  to  sex,  39-43.5  mm.  The  inter- 
mediates measured  29-40  mm.,  except  for  one  25.5  mm.,  average  33  mm., 
mean  34  mm.  What  is  the  transformation  size? 

General  appearance:  "A  localized  derivative  of  Bttfo  boreas,  closely  similar 
to  B.  b.  nelsoni  in  its  small  size,  narrow  head,  and  smoothness  of  skin,  but 
sharply  distinguished  from  nelsoni,  from  all  forms  of  boreas,  and,  indeed, 
from  any  other  North  American  Btifo  by  its  strange  color.  The  dark  dorsal 
markings  of  boreas  have  enlarged,  fused,  and  darkened  until  the  upper  sur- 
faces are  almost  entirely  a  shining  lacquer  black  (deep  dull  blackish  brown 
in  alcohol;  grayish  black  in  formalin),  the  remnants  of  the  light  interspaces 
remaining  as  irregular,  whitish,  or  brownish  vermiform  markings,  and  the 
vertebral  line  showing  as  a  white  or  whitish  hairline,  frequently  greatly  inter- 
rupted, down  the  middle  of  the  back.  The  markings  of  the  underside  are  even 
more  remarkable  and  diagnostic.  The  sparse  black  spots  of  boreas  boreas  or 
boreas  nelsoni  have  developed  into  a  dense  mottling  or  marbling  of  black, 
which  appears  in  life  as  if  made  with  india  ink.  Not  only  are  the  belly  and 
lower  surfaces  of  the  tibiae  and  tarso-metatarsals  heavily  marbled,  but  the 
throat  is  usually  spotted,  and  the  undersides  of  the  femurs  and  rump  are  com- 
pletely black  except  for  the  tubercles,  which  are  white.  The  tarsal  fold  is  very 
poorly  developed  and  is  almost  obsolete  in  most  of  the  specimens"  (G.  S. 
Myers,  Calif.,  1942,  p.  3). 

Color:  Deep  Springs,  Calif.,  May  12,  1942.  Young,  half  grown.  Middorsal 
stripe  is  primrose  yellow  m  one  and  marguerite  yellow  in  another.  The  paro- 
toid  in  one  bears  a  prominent  patch  of  isabella  color,  another  of  olive  ocher. 
This  color  extends  down  the  back  between  tubercles  and  becomes  lighter, 
sometimes  deep  seafoam  green.  The  general  background  of  back  is  light 
brownish  olive  with  about  three  irregular  rows  of  warts,  one  on  either  side  of 
middorsum,  one  back  from  parotoid,  and  an  intermediate  one.  Bordering 
this  light  brownish  olive  area  is  a  broken  irregular  thread  of  marguerite  yel- 
low, below  which  is  a  band  of  almost  clear  black  along  the  sides.  The  black 
of  the  lower  sides  is  broken  into  mottling  by  marguerite  yellow.  Marguerite 
yellow  or  ivory  yellow  is  the  dominant  ventral  color,  very  heavily  spotted  with 
black — so  heavily  spotted  on  rear  of  belly  and  across  thighs  that  the  nickname 
"black  pants"  was  used  in  the  laboratory.  On  the  face  is  an  oblique  band  of 
ivory  yellow  or  marguerite  yellow  extending  backward  from  the  eye,  some- 
times forming  a  half  moon  and  reaching  the  angle  of  the  mouth.  There  is  a 
black  band  from  eye  to  arm  insertion,  bordered  behind  by  a  longitudinal 
stripe  of  marguerite  yellow  or  a  broken  row  of  spots  of  the  same.  The  iris  is 
black  with  prominent  speckings  of  glass  green  and  with  a  wash  of  cinnamon 
in  the  upper  portion;  the  pupil  rim  is  of  reed  yellow.  The  young  are  very  sug- 
gestive of  the  females  of  B.  canorus. 


178  HANDBOOK  OF  FROGS  AND  TOADS 

Female.  The  female  has  very  low  tubercles,  smooth  and  flattened,  the  mid- 
dorsal  stripe  of  the  young,  and  a  few  flecks  of  marguerite  yellow  along  the 
sides  (otherwise  the  dorsum  is  entirely  black).  These  toads  have  the  most 
prominent  marguerite  yellow  and  black  markings  on  the  upper  surfaces  of 
fore  and  hind  limbs,  particularly  on  the  underside  of  tibia  and  dorsum  of  foot. 
The  underside  of  the  foot  is  black,  except  for  the  articular  tubercles  which 
are  light  yellowish  olive.  The  throat  is  more  heavily  spotted  than  in  the  young. 
Iris  as  in  the  young.  The  face  marking  has  the  oblique  light  line,  then  black 
area  from  eye  down,  then  irregular  light  longitudinal  stripe  back  of  angle  of 
mouth. 

Male.  No  striking  difference  from  female.  The  inner  palmar  tubercle  is  very 
prominent. 

This  lack  of  sexual  dimorphism  makes  these  toads  different  from  B.  cano- 
rus,  but  in  general  appearance  they  seem  like  a  smaller  black  B.  canorus. 

Some  random  notes  on  the  Ferris,  Wiggans,  and  Myers  material  follow: 
All  except  one  male  have  thin  light  stripe  full  length  of  body.  One  female  has 
line  absent  in  rear  half  and  broken  in  the  front  half.  One  female  has  line 
broken  in  many  places.  The  young  have  a  prominent  stripe.  The  first  young 
to  have  uniform  black  back  is  40  mm.  long  (no.  2553) ;  those  36  mm.  and  38 
mm.  are  spotted  on  either  side  of  middorsum.  Even  some  of  smallest  have 
black  on  sides.  One  of  striking  things  in  this  form  is  the  thin  light  line  going 
through  interorbital  area  and  the  oblique  bar  crossing  from  one  eyelid  to  the 
other.  In  many  this  cross  thread  is  absent  and  there  is  a  light  longitudinal  spot 
on  parotoid,  but  this  is  absent  in  some.  The  venter  is  heavily  mottled  with 
black  from  pectoral  to  pelvic  region;  the  groin  and  underside  of  femur  and 
pelvic  region  are  black  with  white  pebbling.  On  forward  half  of  venter  the 
white  and  black  are  about  equally  divided.  The  throats  are  lighter,  are  creamy, 
but  with  black  clots  in  most  cases.  Six  of  the  small  ones  have  scant  spotting  on 
throat  region  and  four  with  heavy  dotting.  All  ten  of  these  have  prominent 
black  pectoral  spot.  Some  of  females  look  ripe. 

Structure:  "Description  of  Holotype.  Head  rather  narrow,  its  width  approx- 
imately three  times  in  length  to  vent,  with  a  weak  canthal  angle.  Snout  moder- 
ately rounded  when  viewed  from  above,  the  eyes  projecting  somewhat  beyond, 
the  line  of  the  upper  lip.  In  profile  the  snout  is  bluntly  rounded  but  slop- 
ing and  not  vertically  truncated,  the  upper  lip  being  the  most  anterior  point. 
Nostrils  slightly  nearer  to  eye  than  to  tip  of  upper  lip,  the  distance  from  the 
eye  equal  to  internarial  space  and  to  interocular  space,  which  is  flat.  No  cranial 
crests.  Loreal  region  somewhat  inclined,  slightly  concave.  Depth  of  subocular 
area  equals  half  length  of  exposed  part  of  eye.  Distance  from  nares  to  tip  of 
upper  lip  equals  length  of  exposed  part  of  eye.  Tympanum  indistinct,  its 
upper  posterior  border  partly  obliterated,  vertically  oval  in  form,  its  depth 
equal  to  distance  from  nares  to  eye,  or  not  quite  two-thirds  horizontal  diame- 
ter of  orbit.  Tympanum  close  to  eye,  its  distance  from  latter  scarcely  equal  to 


TOADS:  BUFONIDAE  179 

the  narrower  (horizontal)  diameter  of  tympanum.  Distance  of  tympanum 
from  the  corner  of  the  mouth  (directly  below  it)  equals  slightly  more  than 
.(vertical)  depth  of  tympanum.  Parotoid  glands  moderate,  oval  to  triangular, 
wider  and  more  distinct  posteriorly  than  anteriorly,  where  they  fade  out  just 
before  reaching  eyes,  their  length  approximately  equal  to  distance  from  eye 
to  tip  of  upper  lip  (snout  tip) ;  the  glands  are  much  farther  apart  than  their 
own  width  and  do  not  descend  to  the  sides  appreciably  behind  the  tympanum. 

"Arms  moderately  stout,  the  fingers  entirely  free  of  web.  Third  finger  long- 
est; second  and  fourth  approximately  equal,  reaching  base  of  penultimate 
phalanx  of  third.  First  finger  equal  to  or  barely  surpassing  second.  Subarticu- 
lar  tubercles  single,  well  developed  only  at  base  of  each  finger.  Several  small, 
scattered  tubercles  on  each  palm.  Two  distinct,  enlarged  palmar  tubercles,  the 
outer  large  and  rounded,  its  highest  point  toward  its  distal  end,  the  inner 
about  half  the  size  of  the  outer  and  somewhat  more  convex. 

"Legs  stout  and  relatively  short,  the  tarso-metataisal  joint  reaching  middle 
of  tympanum  when  leg  is  brought  forward,  the  heels  not  quite  touching  when 
femur  and  folded  tibia  are  brought  to  right  angles  with  body.  Toes  more  than 
half  webbed,  the  web  reaching  as  far  as  the  base  of  the  antepenultimate  pha- 
lanx of  the  fourth  toe  and  the  penultimate  phalanges  of  the  other  toes;  the  webs 
are  not  greatly  excised.  Inner  metatarsal  tubercle  moderate,  elongate  oval  in 
form,  its  end  only  slightly  free  and  without  cutting  edge;  the  tarsal  fold  run- 
ning proximally  from  the  tubercle  is  nearly  obsolete  and  only  barely  visible. 
Outer  metatarsal  tubercle  smaller,  rather  conical  and  rounded.  Subarticular 
tubercles  of  toes  single  but  weak"  (G.  S.  Myers,  Calif.,  1942,  pp.  4-5). 

Comparison  of  comparable  Bufo  canonts  and  Bttfo  exsul t  each  at  44  and  56 
mm.,  shows  head  wider,  eye  and  interorbital  space  slightly  larger,  tympanum 
much  larger,  fore  limb  longer  in  B.  canoms,  and  third  toe  slightly  longer  in 
B.  exsul.  One  must  remember  that  the  Bufo  canorns  is  hardly  mature  at  44 
mm.  and  reaches  75  mm.  in  extreme  size,  whereas  B.  exsul  is  mature  at  44  and 
reaches  only  61.5  mm.  in  extreme  size. 

Breeding:  On  May  12,  1942,  two  different  sizes  of  tadpoles  were  with  the 
adults:  many  tadpoles  were  19  or  20  mm.  in  length,  and  a  mature  tadpole  set 
were  29-35  mm.,  with  body  lengths  such  as  12.5,  13.0,  13.5,  13.5,  14.0,  and  15.0 
mm.  for  six  we  measured.  The  tooth  formulae  are  %.  The  tail  tip  is  very 
rounded. 

Journal  notes:  May  12,  1942.  Went  to  Deep  Springs.  At  12:30  arrived  at 
Springs,  5000  feet  elevation  7  Mi  miles  south  of-  Deep  Springs  School  and  above 
Deep  Springs  Lake.  Just  below  the  first  spring,  in  the  water  runways  between 
tussocks,  began  to  see  Bufo  exsul.  The  bottoms  are  mud,  water  2  to  4  inches 
deep.  These  areas  more  or  less  shaded  by  tussocks.  Soil  dark  and  mucky.  De- 
pressions between  tussocks  8  inches  to  i  !/£  feet  deep.  The  toads  often  dashed 
into  holes  in  tussocks  or  into  shaded  runways.  The  first  three  running  springs 
unite  into  one  clear  stream.  In  the  stony  clear  sources  no  toads;  just  above 


180  HANDBOOK  OF  FROGS  AND  TOADS 

confluence  of  three  springs,  where  streams  are  not  so  fast,  among  some  rushes 
and  muck,  several  of  the  biggest  ones  were  found.  One  ran  into  a  hole  under 
a  tussock.  Along  this  stream  4  feet  wide  and  nowhere  more  than  2  feet  deep, 
usually  i  foot  or  less,  were  many  toads.  Every  so  often  toads  would  swim  out 
from  bank  and  stay  poised  on  top.  Often  they  would  swim  downward  and 
hide  in  vegetation.  Occasionally  they  would  rush  into  the  mud  but  seldom 
completely  out  of  sight.  Sometimes  one  could  see  in  one  area  as  many  as  3  or 
4  toads  on  bottom  or  poised  in  water  below  surface.  Often  a  shaded  spot  or 
place  with  no  shade  would  have  2,  3,  4,  or  even  6  toads  in  one  localized  area. 
Are  they  approaching  breeding?  No  mated  pairs.  In  this  area  one  also  could 
see  toads  in  the  vegetation  at  surface  or  see  one  or  two  poised  on  a  mat  of  algae 
at  the  surface.  The  farthest  from  the  stream  we  took  toads  was  6  feet.  Even 
near  stream  they  made  for  stream  and  escaped  in  it. 

In  the  stream  were  three  different  sizes  of  tadpoles.  Some  seemed  mature 
but  no  transformation  sizes  were  available.  The  warm  sulphur  spring  beyond 
had  just  as  many  toads.  Some  of  the  largest  ones  are  blacker,  some  of  the 
smaller  ones  were  somewhat  yellowish.  Mr.  Dieflfenderfer  said  he  and  another 
found  2800  of  them  frozen  in  the  water  of  Antelope  Spring  west  of  Deep 
Springs  Valley  Lake.  Did  the  big  tadpoles  come  from  breeding  in  March  this 
year? 

May  13.  Started  about  8:30  from  Deep  Springs.  Snow  all  about. 

Dr.  Myers  showed  us  his  and  the  Hubbs  material  from  Deep  Springs  pre- 
vious to  the  publication  of  his  new  form  (September,  1942). 

Authorities9  corner: 
G.  S.  Myers,  Calif.,  1942,  pp.  8-9. 

Canadian  Toad,  Winnipeg  Toad,  Manitoba  Toad 

Bttfo  hemiophrys  Cope.  Plate  XXXVI;  Map  10. 

Range:  North  Dakota,  Manitoba,  Alberta  to  northwest  provinces.  In  coun- 
try tributary  to  Lake  Winnipeg  in  Canada.  Some  range  discussions  include 
Minnesota,  South  Dakota,  Montana,  and  Mackenzie. 

Habitat:  Stream  valleys,  lakes,  and  Turtle  Mountain  region. 

Size:  Adults,  2^4-3%  inches.  Males,  56-68  mm.  Females,  56-80  mm. 

General  appearance:  These  are  brownish  or  greenish  toads  of  medium  size 
and  with  short  snouts.  The  most  marked  characteristic  is  the  boss  on  the  head. 
This  is  a  narrow  raised  horny  structure  extending  from  the  rear  of  the  upper 
eyelids  to  the  snout.  Frequently  the  rear  end  of  this  boss  forms  quite  a  promi- 
nence. It  makes  one  think  of  a  frontal  plate  of  the  upper  mandible  of  some 
rail  and  cootlike  birds.  Their  backs  are  covered  with  many  fine,  rounded 
warts,  generally  two  to  five  to  a  dark  area.  There  are  many  spiny  warts  on 
the  femur.  There  is  a  light  stripe  down  the  middle  of  the  back  and  a  conspicu- 
ous dark  band  along  the  side,  bordered  above  by  a  light  line.  The  under  parts 


TOADS:  BUFONIDAE  181 

are  cream  or  buflfy  with  numerous  small  black  spots,  a  prominent  one  in  the 
middle  of  lower  throat.  The  throat  of  the  male  is  dark.  The  broad  upper  eye- 
lids with  closely  set  tubercles  are  in  sharp  contrast  with  the  narrow  inter- 
orbital.  This  makes  the  eyes  stand  out. 

Color:  Walhalla,  N.D.,  Sept.  i,  1930.  Male.  The  stripe  down  the  middle  of 
the  back  is  white  or  ivory  yellow,  edged  on  either  side  with  reed  yellow.  From 
eye  backward  over  lower  edge  of  parotoid  and  along  side  to  groin  extends  a 
cream-buff  line,  becoming  chamois  on  lower  edge  of  parotoid  and  in  vertical 
bar  that  extends  down  to  the  arm.  Below  the  dark  lateral  band,  the  interstices 
among  the  dark  spots  are  the  same  color.  The  color  of  the  back  in  its  lighter 
portions  is  ecru-olive;  in  darker  portions,  it  is  yellowish  olive  or  dark  greenish 
olive.  The  dark  bar  on  either  upper  eyelid,  the  area  just  inside  the  inner  tip 
of  parotoid,  and  the  succeeding  paired  spots  along  median  line  are  dark  green- 
ish olive  or  olive  with  brownish  olive  warts.  The  band  on  side  of  body  is  also 
dark  greenish  olive  or  dark  olive.  The  dark  oblique  bar,  below  eye,  and  big 
bar  on  shoulder  insertion  are  the  same  color.  The  tympanum  and  interstices 
between  these  bars  are  malachite  green.  The  arms  and  legs  arc  barred  with  the 
dark  color  of  the  dorsal  spots,  with  interstices  the  color  of  back  and  cream 
color.  The  rear  of  femur  has  the  dorsal  background  color  with  strontian  yel- 
low or  wax  yellow  spots.  This  color  comes  into  the  color  of  the  lower  belly  and 
underside  of  femur.  The  belly  is  cream-buff,  heavily  marked  with  light  gray- 
ish olive.  In  middle  of  pectoral  region  and  on  lower  throat  is  long  bar  spot  of 
the  same.  The  throat  is  suffused  with  light  grayish  olive.  Upper  portion  of 
parotoid  is  drab.  The  boss  is  argus  brown.  The  inner  metatarsal  tubercle  is 
mummy  brown.  The  upper  surfaces  of  first  two  fingers  are  covered  with  ex- 
crescences of  same  color  or  of  chestnut-brown.  The  eye  is  olivaceous  black  or 
dark  ivy  green  with  prominent  lemon  yellow  ins  ring;  upper  part  of  iris  is 
heavily  spotted  with  same  color,  lower  slightly  spotted,  and  upper  and  lower 
parts  slightly  spotted  with  russet-vmaceous. 

Female,  young.  The  spots  on  belly  are  larger;  the  throat  is  clear  of  spots  ex- 
cept for  pectoral  spot.  Throat  is  cream-colored.  First  two  fingers  without  ex- 
crescences. Otherwise  marked  as  in  the  male.  The  outer  circle  of  the  iris  bears 
a  deep  bluish  glaucous  ring. 

(From  preserved  material  in  USNM.)  Back  brown  with  a  median  light 
band.  A  broad  brown  lateral  band  starts  back  of  the  arm  insertion  extending 
backward  to  the  groin  and  forward  as  a  broken  band  to  the  eye,  above  the 
tympanum.  This  is  bordered  above  by  a  light  margin,  which  sends  a  series  of 
more  or  less  vertical  lines  to  the  arm  insertion,  the  mam  branch  going  forward 
involving  the  parotoid.  This  dark  band  borders  the  light  color  of  the  belly, 
which  is  strongly  mottled  with  dark  along  the  sides.  The  median  line  is  bor- 
dered on  either  side  by  a  dark  area  more  or  less  continuous  to  the  vent.  In 
some  of  the  older  frogs  this  breaks  up  into  detached  spots  about  some  of  the 
collections  of  warts.  The  warts  are  lighter  colored.  Between  the  dorsal  series 


182  HANDBOOK  OK  FROGS  AND  TOADS 

of  brown  spots  and  the  lateral  band  are  two  rows  of  spots,  the  more  regular 
one  being  above  the  light  margin  of  the  dark  lateral  band.  There  is  a  brown 
area  back  of  the  eye  and  below  the  tympanum,  a  brown  suborbital  spot,  and 
a  bar  from  nostril  downward;  between  this  bar  and  suborbital  spot  is  another 
bar.  Therefore  the  upper  lip  is  an  alternation  of  light  snout,  nasal  bar,  light 
interval,  dark  bar,  light  interval,  suborbital  dark  spot,  light  interval,  and  sub- 
tympanic  spot.  There  is  a  light  line  from  below  the  tympanum  or  near  angle 
of  the  mouth,  backward  over  the  arm  insertion;  it,  with  the  vertical  bar  from 
the  light  side  band,  extends  to  the  dorsum  of  the  arm.  The  upper  eyelid  has 
a  prominent  brown  transverse  band,  margined  in  front  and  below  by  a  con- 
spicuous light  gray  margin.  This  eyelid  coloration  stands  out  in  sharp  contrast 
to  the  boss  of  the  head.  Breast  and  belly  spotted  with  dark.  The  hind  legs  have 
dark  cross  bands  with  reddish  tubercles  in  the  center  of  cross  bands.  Dark 
reticulations  prominent  on  back  of  femur. 

Structure:  Boss  on  head  from  snout  to  rear  of  interorbital  space;  slight  fur- 
row down  the  middle  and  outer  ridges  forming  boss  are  parallel;  these  ridges 
connected  by  a  bar  at  rear  and  with  slight  extension  toward  the  eye;  broad 
upper  eyelid  with  closely  set  tubercles,  in  sharp  contrast  with  narrow  inter- 
orbital;  snout  short,  vertical  in  profile;  tympanum  elliptical  and  vertical;  two 
cutting  metatarsal  tubercles,  inner  large  and  spadelike;  ventro-basal  portion 
of  femur  discolored;  fore  throat  of  male  discolored. 

Voice:  "Its  spring  note  is  a  soft  trilling,  uttered  about  twice  a  minute  and 
lasting  about  three  seconds"  (E.  T.  Seton,  Man.,  1918,  p.  83). 

Breeding:  This  species  breeds  from  May  onward,  in  shallow  edges  of  lakes 
or  ponds  or  other  water.  There  are  no  descriptions  of  the  tadpole  on  record. 
They  transform  at  %-"!/>  inch  (9.0-13.5  mm.). 

Journal  notes:  August  30,  19 30.  At  n  o'clock  we  started  northward  from 
Grand  Forks.  At  Ardock  we  found  a  stream  dried  up  except  for  isolated 
pools.  Around  one  in  the  cow-punched  and  cracked  mud,  we  took  some  seven 
or  eight  young  of  Btijo  hemiophrys.  They  were  of  two  sets:  36  mm.,  33  mm., 
26  mm.,  23  mm.,  22  mm.,  and  18  mm.  All  of  these  young  specimens  have  a 
suggestion  of  the  dark  pectoral  spot,  and  only  the  two  larger  ones  begin  to 
show  cranial  ridges. 

August  31,  1930.  It  rained  in  the  afternoon.  We  went  through  Neche  to 
Walhalla,  N.D.,  at  the  edge  of  Pembina  Mountains.  We  arrived  about  5:30 
P.M.  It  was  still  raining.  We  began  to  get  desperate  in  our  desire  for  live  adults 
of  Biifo  hemiophrys.  Before  dark  we  took  an  excursion  around  town  looking 
for  possible  covers  for  toads.  The  evening  was  cold,  the  rain  had  stopped,  and 
it  was  doubtful  if  toads  were  abroad  feeding.  This  has  been  a  summer  of  re- 
markable drought.  We  used  one  of  the  best  expedients  of  a  collector.  We  en- 
gaged a  bright-eyed  young  boy  (Audrey  Miller)  to  search  the  town.  Inside 
of  half  an  hour  he  appeared  with  an  adult  male.  He  and  another  lad  started 
promptly  again  with  a  flashlight  and  returned  twice,  each  time  with  an  adult 


TOADS:  BUFONIDAE 


183 


Plate  XXXVI.  Bufo  hcmiophrys.  i.  Male  Plate  XXXVll    Bufo  msid.or   (X%)« 

(X%).  2,4,5.   Males   (X%)-  3-  Young         Males,  from  Toyahvale,  Tex. 

/i      .n  /  \ 


184  HANDBOOK  OF  FROGS  AND  TOADS 

toad.  We  asked  where  they  got  them  and  he  said  in  the  gutters,  and  later  he 
said  he  got  them  in  the  sewers. 

Sept.  i,  1930.  We  went  out  with  the  boys  to  a  place  where  they  had  a  toad 
located,  but  had  missed  him  last  night.  We  wondered  why  they  carried  a  flash- 
light and  a  pail  on  a  long  rope.  We  soon  found  out.  The  town  has  drainage 
manholes.  The  boys  took  off  the  cover  and  with  flashlight  searched  the  surface 
of  the  water  and  the  walls  of  the  hole.  They  pointed  out  a  toad  floating  at  sur- 
face. They  let  down  their  pail  and  drew  it  up  under  the  toad  and  so  caught  it. 
They  brought  up  two  toads  instead  of  one,  but  one  was  dead.  Quite  likely  the 
toads  were  caught  here  as  in  a  trap,  but  at  this  long  period  of  drought  it  was 
our  best  chance  of  getting  material. 

These  males  have  a  throat  of  the  Bufo  americantts  class,  but  close  examina- 
tion of  one  which  half  inflated  its  throat  revealed  that  the  lower  throat  region 
expanded  first,  and  there  also  seemed  to  be  a  secondary  swelling  at  the  ventral 
base  of  each  arm.  The  throat  seems  to  have  a  slight  fold  across  the  lower  throat 
on  the  pectoral  region.  Its  boss,  this  slight  indication  of  a  lower  throat  pouch, 
and  two  cutting  mctatarsal  tubercles  suggest  relationship  with  B.  cognattts. 

Authorities'  corner: 
E.  T,  Seton,  Man.,  1918,  p.  83. 

Sonoran  Toad 

Bufo  tnsidior  Girard.  Plate  XXXVII;  Map  12. 

Range:  Colorado  and  Kansas  south  through  New  Mexico,  Arizona,  and 
Texas  into  northern  Mexico. 

Habitat  and  Size:  Since  Dr.  Taylor's  (1936)  resurrection  of  this  Girard  spe- 
cies from  the  synonymy  of  Bufo  debihs  and  since  the  inclusion  of  it  in  the 
check  lists  (Stejnegcr  and  Harbour),  little  material  has  appeared  to  add  to 
our  understanding  of  it. 

General  appearance:  From  our  limited  experience  with  this  form  in  Trans- 
Pecos,  Tex.,  we  would  call  it  more  bleached  than  are  southern  Texan  little 
green  toads. 

"The  absence  of  cranial  crests,  or  the  presence  of  very  narrow  ones  closely 
approximated  to  orbits,  coupled  with  the  broad,  elongate  parotoid  gland  and 
greenish  coloration  is  very  distinctive  of  this  Kansas  toad"  (H.  M.  Smith, 
Kans.,  1934,  p.  443). 

Color:  July  22,  1942,  Toyahvale,  Tex.  This  toad  has  a  flat  head,  pointed 
snout,  and  very  large,  elongate,  and  broad  parotoids.  The  back  of  one  is 
courge  green,  with  the  top  of  the  head  clear  yellow-green.  Another  has  a  bis- 
cay  green  back  with  top  of  snout  and  upper  eyelids  touched  with  old  gold. 
The  back  and  limbs  bear  tubercles  encircled  with  mustard  yellow  and  tipped 
with  mummy  brown.  The  back  and  top  of  head  are  also  flecked  with  black, 


TOADS:  BUFONIDAE  185 

each  spot  with  its  black  central  tubercle.  Some  spots  are  dumbbells  in  shape. 
A  prominent  oblique  black  bar  crosses  the  eyelid.  The  hind  limbs  have  two 
or  three  bars  on  femur,  two  on  tibia,  and  one  on  metatarsus.  On  the  sides  the 
tubercles,  encircled  with  chartreuse  yellow  and  tipped  with  black  or  mummy 
brown,  are  on  a  mummy  brown  background,  which  forms  a  broken  line  across 
the  rear  of  parotoid  and  along  the  side.  The  groin,  underside  of  thighs,  and 
sides  of  belly  are  pale  brownish  vinaceous;  the  center  of  the  belly  is  light  glau- 
cous-blue, one  immaculate,  another  with  several  spots.  The  throat  of  the  fe- 
male is  white,  that  of  the  male  discolored,  the  forward  part  deep  purplish 
vinaceous,  the  rear  a  pale  vinaceous-lilac,  the  middle  deep  medici  blue.  The 
underside  of  hand  and  of  toes  is  vinaceous-cinnamon,  toes  and  fingers  tipped 
with  cinnamon-brown.  The  two  metatarsal  tubercles  and  the  palmar  tubercle 
are  russet.  The  black  eye  has  a  citron  yellow  pupil  rim  broken  in  four  places, 
and  the  iris  is  heavily  flecked  with  ochraccous-buff. 

Structure:  The  original  description  (C.  Girard,  Gen.,  1856,  p.  88)  is  as  fol- 
lows: "Upper  surface  of  head  plane  and  smooth.  Snout  subacute,  protruding. 
Mouth  moderate,  upper  jaw  slightly  emarginated.  Tongue  elongated  tapering 
towards  both  ends.  Tympanum  inconspicuous.  Parotoids  large  and  elongated, 
situated  obliquely  upon  the  shoulder.  Limbs  moderate.  First  finger  equal  to 
the  second  in  length.  A  metacarpal  disk,  and  a  tubercle.  Toes  slightly  webbed 
at  base.  Two  metatarsal  tubercles.  Skin  papillous  above,  warty  beneath." 

Voice:  "This  species  is  rather  abundant  throughout  the  region  traversed 
(eastern  border  of  the  Staked  Plain  of  Texas) .  It  is  frequently  found  in  the 
grass,  where  ils  green  color  aids  in  concealing  it.  When  in  the  water,  its  cry 
is  like  that  of  B.  lentiginosus  americantts,  but  is  more  feeble,  and  very  nasal" 
(E.  D.  Cope,  Tex.,  1893,  p.  332). 

Breeding:  Like  B.  debtlis,  it  doubtless  breeds  from  mid-March  to  August. 
Ripe  females  have  been  secured  to  mid-July. 

Journal  notes:  Since  the  re-establishment  of  Bttfo  insidior,  we  have  had  no 
opportunity  to  examine  the  collections  of  Bujo  debilis  and  Bujo  insidior  to 
outline  their  contrasting  characters  or  to  determine  their  actual  distinctness. 
The  instant  we  saw  Taylor's  range  of  /?///<?  insidior  we  thought  of  the  time, 
July  10,  1925,  near  Fort  Davis,  Tex.,  when  we  heard  narrow-mouthed  toads 
and  green  toads;  and  also  of  the  time  in  1934  when  we  heard  these  same  little 
toads  at  Alpine,  Tex.  On  our  1942  trip  we  secured  the  toad  near  Balmorhca 
State  Park. 

July  22,  1942,  Toyahvale.  Night  trip  west  on  highway.  ...  In  this  pond 
heard  one  lone  Microhyla  olivacea.  Farther  on  heard  plenty  of  Microhyla. 
Then  I  began  to  question  whether  they  were  all  Microhyla.  There  was  a  call 
halfway  between  that  of  a  Microhyla  and  a  Bujo.  A  little  toad  under  my  feet 
put  into  the  pond.  I  didn't  see  it  well.  Then  I  realized  that  it  must  be  old  Bujo 
debilis.  Now  that  we  were  in  the  range  of  Taylor's  Bujo  insidior,  might  it  be 


1 86  HANDBOOK  OF  FROGS  AND  TOADS 

that  form?  Caught  three  males  (36,  58,  and  43  mm.).  As  we  came  back  heard 
many  more,  and  where  we  heard  it  most,  there  on  edge  of  road,  saw  a  female 
just  hit  but  yet  alive. 

When  we  compared  three  Toyahvale  specimens  of  B.  insidior  with  three 
of  our  southern  Texas  B.  debilis  we  seemingly  had  these  differences: 

B.  insidior  B.  debilis 

Snout  6.9-7.8  in  L.  8.0-9.2  in  L. 

Upper  eyelid  9.5-10.0  in  L.  11.5-12.5  in  L. 

ist  finger  7.6-8.6  in  L.  8.8-11.5  in  L. 

3rd  finger  4-4-4-5 in  L.  5.2-6.5  in  L. 

4th  finger  5.06-5.5  in  L.  6.5-8.0  in  L. 

Foot  with  tarsus  1.7-1.8  in  L.  2.0-2.15  in  L. 

Whether  these  measurements  will  obtain  with  a  large  series  we  dare  not  as- 
sert. They  point  to  bigger  snout,  upper  eyelid,  first,  second,  and  fourth  fingers, 
and  longer  foot  with  tarsus  in  B.  tnstdwr. 

Authorities'  corner:  "In  September,  1886,  coming  down  the  valley  of  the 
Cimarron  river  from  New  Mexico,  I  first  noticed  the  little  Sonoran  toad,  Bufo 
debilis,  Girard,  near  the  Z  H  ranch  in  the  Public  Lands  (now  Beaver  county, 
Oklahoma),  at  a  point  thirty-five  or  forty  miles  west  of  the  southwest  corner 
of  Kansas.  The  species  was  observed  a  few  days  later  in  great  abundance  and 
activity  (during  rainy  weather)  in  Morton  county,  Kansas,  and  in  the  south- 
ern part  of  Hamilton  county.  I  have  collected  a  single  specimen  in  the  western 
part  of  Barber  county,  Kansas,  also"  (F.  W.  Cragin,  Kans.,  1894,  p.  39). 

"Two  specimens  (U.S.N.M.  no.  2622)  from  Chihuahua,  collected  by  Dr. 
Thomas  H.  Webb,  are  designated  in  the  museum  catalogue  as  the  cotypes  of 
Girard's  Btifo  insidior.  The  preservation  of  these  specimens  is  fair,  but  both 
are  very  much  bleached"  (R.  Kellogg,  Gen.,  1932,  p.  52). 

When  Dr.  E.  H.  Taylor  described  a  new  species  from  a  locality  near  Mazat- 
lan,  he  said  that  this  small  toad,  Bufo  folloggi  (named  in  honor  of  Dr.  Rem- 
ington Kellogg)  "is  most  closely  related  to  Bufo  debilis  and  Bufo  insidior 
Girard."  "From  Btifo  insidior  Girard  (Kansas,  Texas,  New  Mexico,  Chihua- 
hua, Durango,  and  Zacatccas)  it  differs  in  the  presence  of  the  cranial  crests 
(lacking  or  with  only  an  occasional  faint  trace  of  crests  in  insidior),  in  having 
shorter  hind  legs,  a  shorter  snout,  larger  eye,  narrower  head,  and  narrower 
interorbital  width,  a  differently  shaped  parotoid  and  a  totally  different  dorsal 
color  pattern.  It  has  very  much  larger  and  more  numerous  spiny  tubercles  on 
dorsal  and  lateral  surfaces;  the  inner  palmar  tubercle  is  less  developed,  as  are 
the  metatarsal  tubercles;  the  webbing  between  the  toes  is  slightly  more  exten- 
sive. The  color  pattern  is  entirely  different"  (E.  H.  Taylor,  Gen.,  i936a, 
p.  513).  We  have  seen  two  B.  f(elloggi  and  hold  them  distinctly  different. 

"Bufo  insidior  Girard.  The  little  green  toad,  although  common  in  Texas 
and  known  from  Kansas,  has  apparently  not  been  recorded  previously  from 


TOADS:  BUFONIDAE  187 

Oklahoma.  Charles  C.  Smith  and  I  found  it  to  be  fairly  common  in  the 
southwestern  part  of  Oklahoma  where  it  was  breeding  with  B.  compactilis. 
Whereas  B.  compactilis  was  also  found  in  great  numbers  on  roadways,  B.  in- 
sidior  was  seen  only  in  breeding  pools.  We  collected  specimens  in  Cotton, 
Jackson,  southern  Kiowa,  and  Tillman  counties.  In  addition,  Mr.  Kuntz  has 
furnished  one  specimen  taken  on  a  street  during  an  evening  shower  in  Law- 
ton,  Comanche  County;  and  the  University  of  Oklahoma  Museum  of  Zoology 
has  a  single  specimen  from  extreme  northern  Garvin  County"  (A.  N.  Bragg, 
Okla.,  i94ia,  pp.  51-52). 

"Under  the  name  Bufo  debihs,  Campbell  [Ariz.,  1934,  p.  3]  recorded  this 
species  from  the  Huachuca  Mountains  in  Arizona,  and  more  recently  Kauf- 
feld  .  .  .  has  reported  a  specimen  from  the  vicinity  of  Tombstone  (30  miles 
south  of  Cochise).  In  view  of  the  fact  that  Taylor  .  .  .  has  reported  speci- 
mens from  Zacatecas,  it  is  reasonable  to  suppose  that  the  species  occurs  in 
northeastern  Sonora. 

"The  specimen  collected  by  Kauffeld  is  now  A.M.N.H.  No.  50914  in  the 
collection  of  the  American  Museum  of  Natural  History.  This  specimen,  an 
adult  female  with  mature  eggs  in  the  oviducts,  compares  favorably  in  colora- 
tion and  pattern  with  Zacatecan  specimens  depicted  by  Taylor  .  .  .  ,  particu- 
larly the  individual  depicted  by  him  as  figure  9  of  plate  45.  Taylor  states  that 
B.  debilis,  with  which  Kellogg  (1932)  synonymized  insidior,  inhabits  Eastern 
Texas  and  Tamaulipas,  whereas  insidior  occupies  an  extensive  range  from 
Kansas  and  New  Mexico  southward  through  Texas  and  the  adjoining  states 
in  Mexico  to  Durango  and  Zacatecas.  We  have  compared  the  Arizonan  speci- 
men with  individuals  from  Archer,  Presidio,  McLennan,  and  Bexar  counties, 
Texas.  Presumably  some  of  these  specimens  would  be  referable  to  debilts,  if 
Taylor's  views  are  correct.  Obviously  there  are  differences  in  pattern  and  pos- 
tulation,  but  there  is  no  conspicuous  difference  aside  from  these  between  the 
Arizonan  and  Texan  specimens.  The  specimens  from  Archer  County  in 
northern  Texas  are  certainly  the  same  species  as  that  from  the  other  Texan 
counties,  including  Bexar,  which  lies  immediately  north  of  Atascosa  County, 
where  the  toads  depicted  as  debilis  in  figures  5  and  6  of  Taylor's  plate  45  were 
taken.  If  insidior  can  be  removed  from  the  synonymy  of  debilis,  we  strongly 
suspect  that  it  represents  a  subspecies  of  the  latter.  Furthermore,  if  our  speci- 
men from  Bexar  County  is  properly  assigned  to  debtlis  it  is  obvious  that  the 
form  has  a  wider  range  than  Taylor's  statement  implies,  probably  including 
Oklahoma' and  Kansas,  to  judge  only  by  a  photograph  of  a  Kansan  specimen, 
and  the  proximity  of  Archer  County  to  Oklahoma.  San  Diego  County,  Texas, 
where  the  toad  in  figure  4  of  Taylor's  plate  was  reputedly  taken,  seems  to  be 
non-existent;  possibly  the  specimen  came  from  San  Diego,  in  Duval  County. 

"Unless  the  type  or  other  specimens  from  Chihuahua  more  closely  resemble 
the  Arizonan  and  Zacatecan  individuals  than  Texan  specimens,  the  applica- 
bility of  the  name  insidior  to  the  former  is  open  to  considerable  question. 


1 88  HANDBOOK  OF  FROGS  AND  TOADS 

Taylor  does  not  compare  debilis  with  insidior,  but  compares  each  separately 
with  kelloggi  from  Mazatlan.  The  latter  evidently  is  a  valid  form  inhabiting 
the  coastal  plain  from  Mazatlan  at  least  as  far  south  as  Acaponeta  in  Nayarit 
where  the  specimens  (A.M.N.H.  Nos.  43877-43878)  were  secured  by  one  of 
us  in  November,  1939"  (C.  M.  Bogert  and  J.  A.  Oliver,  Gen.,  1945,  pp.  409- 
410).  See  page  87  for  Taylor  and  Smith's  recent  (1948)  comparisons. 

Marine  Toad,  Giant  Toad 

Bufo  marinus  (Linne).  Plate  XXXVIII;  Map  10. 

Range:  From  southern  Sonora  to  Tamaulipas  in  Mexico  to  Patagonia.  In- 
troduced into  Puerto  Rico,  Jamaica,  and  several  smaller  West  Indian  islands, 
some  of  which  may  possibly  have  had  it  originally  native.  Some  distance  north 
of  William  Lloyd's  collection  of  it  in  Hidalgo,  Tamaulipas,  E.  H.  Taylor  and 
J.  S.  Wright  took  it  near  Zapata,  Tex.  They  wrote:  "While  collecting  along 
the  highway  between  Zapata  and  Arroyo  Loma  Blanca,  in  southern  Texas,  on 
the  night  of  August  20,  1931,  encountered  a  number  of  toads  in  a  temporary 
pool,  which  on  examination  proved  to  be  the  widespread  Bufo  marinus 
(Linn.),  the  largest  known  species  of  Bufo." 

Habitat:  In  1891  Ives  found  it  in  the  courtyard  of  a  house.  In  1901  Waite  in 
Bermuda  reported  them  around  houses,  in  gardens,  and  in  water  tanks.  In 
1917  in  Bermuda  P.  H.  Pope  stated  that  they  were  "abundant  about  roadsides, 
gardens  and  edge  of  mangrove  swamps.  A  street  light  halfway  between  'Gras- 
mere'  and  Hamilton  was  a  favorite  place  for  them,  and  two  or  three  were 
usually  seen  under  it,  picking  up  the  insects  that  were  attracted  by  the  light. 
...  In  the  day-time  they  hide  under  stones  or  boards  or  burrow  in  the  soft 
earth.  I  have  sometimes  seen  them  in  little  burrows  in  the  side  of  a  bank, 
where  they  had  dug  themselves  in  just  enough  to  be  out  of  the  sun."  In  1932, 
R.  Kellogg  declared  that  "these  large  toads  are  nocturnal  in  their  habits  and 
hide  under  fallen  tree  trunks,  matted  leaves,  and  stones,  or  burrow  into  loose 
soil." 

Size:  One  synonym,  Bufo  gigas  Walbaum,  suggests  its  size.  It  is  the  giant 
of  toads.  Gunther  in  1858  (Gen.)  remarked  on  the  very  large  specimens  and 
in  1885-1902  he  noted  that  "Central- American  specimens,  at  least  those  of  the 
more  temperate  districts,  do  not  appear  to  grow  to  the  enormous  size  to  which 
the  species  attains  in  Brazil  and  Guiana."  His  measured  specimen  is  166  mm. 
P.  H.  Pope  (1917)  in  Bermuda  wrote,  "An  average  female  measured  145  mm. 
from  snout  to  vent  and  Waite  describes  one  155  mm.  in  length."  "The  male 
is  smaller,  usually  about  13  mm.  shorter,  and  more  active  than  the  female." 
Schmidt  in  1928  had  a  i03-mm.  female  and  a  95-mm.  female  from  Puerto  Rico. 
Kellogg  (1932),  among  the  104  specimens  he  examined,  mentioned  individ- 
uals measuring  no  mm.,  125  mm.,  and  167  mm.  Taylor  and  Wright  (1932) 
took  several  measuring  from  66-168  mm.,  but  the  record  seems  to  be  that  of 


TOADS:  BUFONIDAE  189 

Miranda-Ribero,  who  reports  a  body  length  of  22  cm.  (220  mm.)  or  8.6  inches. 

General  appearance:  This  is  an  enormous  toad  with  an  immense  triangular 
pitted  parotoid.  The  forward  side  of  this  triangular  gland  is  just  back  of  the 
tympanum;  the  base  starts  just  back  of  the  angle  of  the  jaw  and  slants  slightly 
upward  to  meet  the  very  oblique  or  slanting  upper  side  at  a  point  well  back  on 
the  side  of  the  body.  From  this  point  a  row  or  fold  of  large,  blunt,  rounded, 
brown-tipped  warts  extends  %  to  %  of  the  way  to  the  groin.  From  Bitfo  valli- 
ceps  it  may  be  distinguished  by  its  very  large  parotoids;  larger  outer  sole 
(metatarsal)  tubercle;  its  coarser,  less  even,  and  less  pointed  tubercles;  blunter 
and  lower  cranial  crests;  the  usual  absence  or  indistinctness  of  the  parietal 
ridge;  and  less  distinct  color  pattern. 

Color:  We  have  seen  none  alive  in  recent  years  and  consequently  use  Dr. 
Kellogg's  (Gen.,  1932)  summary:  "In  general,  the  coloration  of  Bitfo  marintts 
is  quite  variable,  ranging  through  various  shades  of  brown,  including  yellow- 
ish, reddish,  and  even  blackish,  and  occasionally  greenish  olive;  upper  parts 
with  or  without  large  msuhform  spots,  which  when  present  are  usually  edged 
with  pale  yellow;  a  light  vertebral  line  occasionally  visible;  arms  and  legs  of 
immature  individuals  usually  banded  with  dark  brown;  underparts  dingy 
white  or  yellow." 

Structure:  Interorbital  region  smooth,  decidedly  concave,  shallower  in  front 
than  behind;  crests  pronounced — the  canthus  rostrahs  is  a  prominent  crest 
beginning  ahead  of  and  above  the  nostrils  and  ending  at  the  anterior  corner 
of  the  eye,  where  it  forks  into  two  ridges,  a  broad  preorbital  and  a  well-defined 
supracihary  crest  which  curves  around  the  eye,  sending  off  a  prominent  supra- 
tympanic  ridge  to  the  parotoid  and  a  very  short  postorbital;  parietal  crests  ab- 
sent, indistinct,  or  poorly  developed;  large  broad  head,  box-snouted;  parotoids 
huge,  as  long  as  head  or  bigger,  %  to  i  l/*  times  the  head;  parotoids  widely  di- 
vergent behind;  toes  M»  to  %  webbed;  free  inner  metatarsal  tubercle;  outer 
metatarsal  tubercle  large  and  flat;  a  thin  edged  tarsal  fold  from  inner  meta- 
tarsal tubercle  backward;  prominent  palmar  pads  at  base  of  first  and  second 
fingers;  eyelid  finely  tuberculate;  two  rows  of  large  fleshy  warts,  one  down 
each  side  of  middorsum  to  vent,  more  prominent  in  male.  Back  of  angle  of 
mouth  the  skin  is  divided  into  two  or  more  vertical  folds  so  that  at  times  it 
might  be  thought  a  postrictal  gland.  However,  it  is  not  like  the  white  wart  of 
Bufo  alvarius. 

Voice:  Taylor  and  Wright  heard  these  toads  at  Zapata,  Tex.,  but  did  not 
describe  their  call.  Pope  in  Bermuda  has  stated  that  they  have  a  more  resonant 
and  louder  call  than  Bufo  amertcanus — "a  deep  booming  trill." 

Breeding:  In  Bermuda  February  to  July  is  the  breeding  season,  April  being 
the  optimum  month.  In  Trinidad  they  breed  from  August  to  October;  in 
Demerara,  from  mid-April  to  September,  possibly  also  from  November  to 
January.  The  eggs  are  in  strings.  Dr.  E.  L.  Mark  had  some  hatch  in  68  hours. 
The  black  tadpole  transforms  after  45  days  or  less.  The  tops  of  the  first  two 


190  HANDBOOK  OF  FROGS  AND  TOADS 

fingers  of  the  males  have  excrescences,  and  there  are  excrescences  on  the  inner 
side  of  third  finger  and  the  inner  palmar  tubercle.  Some  males  have  brown 
and  spiny-tipped  warts  and  tubercles. 

Clark  has  a  note  that  this  species  lays  twice  a  year,  but  Pope  is  sure  that  in 
Bermuda  an  individual  breeds  but  once  a  year.  In  Bermuda  in  1903  Mark 
made  the  following  observations : 

"In  the  spring  of  1903  ...  we  visited  'Spanish  Rock'  near  midday,  April 
22.  Before  reaching  the  Rock  we  found,  on  the  slope  facing  away  from  the 
ocean  and  toward  Spittal  Pond,  a  small  pool  of  rainwater  (there  had  been  a 
heavy  shower  the  night  before),  and  in  this  pool  there  were  large  numbers 
of  the  huge  toads  Bit  jo  aqua — in  pairs.  The  females  were  engaged  in  spawn- 
ing, and  the  numerous  strings  of  spawn  were  stretched  across  the  pool  in 
almost  every  conceivable  direction.  The  pool,  some  fifteen  or  twenty  feet  in 
diameter  and  only  a  few  inches  deep,  was  of  so  temporary  a  character  that  its 
bottom  consisted  throughout  of  turf,  not  unlike  the  land  immediately  sur- 
rounding it,  which  was  not  submerged." 

Ruthven  (1916)  "recorded  that  on  the  Demerara  River,  about  thirty-five 
miles  south  of  Georgetown  m  1914,  tadpoles  were  abundant  in  July  and  Au- 
gust. .  .  ."  Pope  (1917)  thought  that  development  might  be  more  rapid  than 
Ruthven's  45  days  from  egg  to  adult.  Mark  (1903)  observed  this  species  from 
eggs  April  22  to  free-swimming  larvae  April  27.  In  describing  some  tadpoles 
collected  by  L.  J.  Cole  in  Bermuda  Pope  wrote:  "The  tadpoles  are  black  in 
color  and  resemble  those  of  B.  americantts.  They  measure  from  8.5  to  10.9  mm. 
in  length.  They  have  the  typical  early  tadpole  form.  The  buds  of  the  hind 
limbs  are  present,  but  hardly  visible  to  the  naked  eye." 

Transformation  must  be  over  as  extended  a  period  as  is  the  egg  laying. 
Ruthven  records  it  on  August  16  about  45  or  46  days  after  egg  laying.  No 
other  notes  apparently  exist  on  transformation. 

Journal  notes:  We  present  none.  Our  experiences  consist  in  seeing  a  few 
captive  specimens  in  aquaria  or  zoos. 

Authorities9  corner:  Taylor  and  Wright  (Tex.,  1952,  pp.  247-249)  found 
their  specimens  had  been  feeding  on  "tenebriomd  and  carabid  beetles,  the 
bulk  of  the  food  being  of  the  genera  Eleodes  and  Pasimachus."  Pope  (1917) 
says  that  when  the  residents  of  Bermuda  found  that  these  toads  ate  cock- 
roaches, they  no  longer  called  them  after  their  introducer  "Captain  Vesey's 
nuisances."  In  1918  Noble  found  that  the  Bufo  marinus  he  had  from  Nica- 
ragua fed  principally  on  cockroaches.  Our  friends,  Drs.  M.  L.  Leonard  and 
Stuart  Danforth,  years  ago  (1930  or  earlier)  told  us  how  some  of  their  co- 
laborers  in  Puerto  Rico  had  successfully  shipped  them  to  Hawaii  to  control 
insects,  and  today  plenty  of  reports  appear  on  their  role  in  Hawaii. 

Regarding  the  poison  Vernll  says:  "This  toad  is  believed  ...  to  have  a 
very  poisonous  secretion  from  its  parotoid  and  dorsal  glands.  It  is  said  that 


TOADS:  BUFONIDAE  191 

dogs  that  mouth  them  invariably  die  within  a  few  hours.  The  secretion  of  the 
glands,  when  injected  into  the  circulation  of  dogs,  birds,  and  other  animals, 
causes  convulsions  and  death,  even  when  in  small  doses.  Mr.  A.  H.  Verrill 
...  on  one  occasion  saw  the  venom  ejected  as  a  fine  spray,  from  the  parotoid 
glands  of  a  large  toad,  when  it  was  much  irritated." 

On  this  topic  the  observations  of  C.  T.  Dodds  seem  pertinent:  "During 
the  summer  of  1922  I  spent  some  time  at  Los  Mochis,  in  Northern  Sinaloa, 
Mexico.  It  was  very  interesting  each  evening  to  note  the  appearance  of 
large  toads  under  the  electric  lights.  One  evening,  about  the  end  of  July 
while  one  of  these  toads  was  occupied  in  catching  insects,  a  small  terrier  dog 
started  to  tease  the  amphibian.  At  first  the  toad  only  hopped  in  an  effort  to 
escape  the  barking  dog,  which  did  not  attempt  to  bite.  Soon  the  toad  became 
tired  and  its  hops  were  less  frequent  and  not  so  long.  The  dog  was  urged  on 
by  the  bystanders  and  finally  snapped  the  toad  while  the  latter  was  in  mid- 
air, catching  it  just  back  of  the  foreleg.  Although  the  dog's  mouth  was  not  in 
contact  with  the  toad  for  more  than  an  instant,  he  immediately  lost  all  inter- 
est in  the  animal.  Spitting  and  shaking  his  head,  he  gave  all  indications  of 
having  received  something  very  distasteful  m  his  mouth.  He  was  offered 
some  water  but  refused  to  drink  and  in  about  a  minute's  time  showed  signs 
of  weakening  and  general  paralysis.  He  sank  to  the  ground  with  his  legs 
spread  out,  writhing  and  whining  with  pain,  and  unable  to  recognize  his 
master.  During  this  time  he  was  able  to  push  himself  along  the  ground,  grad- 
ually becoming  weaker  and  very  rigid,  with  eyes  greatly  protruding  and 
respiration  and  heart  action  exceedingly  rapid.  After  twenty  minutes  he  was 
somewhat  quieter  as  if  he  was  going  to  die.  It  was  suggested  that  castor  oil 
be  given  him  in  the  hope  that  it  might  save  his  life.  Accordingly  his  mouth 
was  pried  open,  for  every  muscle  was  rigid,  and  about  50  cc.  of  oil  was  poured 
down  his  throat.  Ten  minutes  later  he  showed  signs  of  improvement,  al- 
though I  do  not  know  that  the  oil  had  anything  to  do  with  the  change.  Within 
another  15  minutes  he  was  able  to  get  upon  his  feet  but  was  very  feeble  and 
his  hind  quarters  were  still  somewhat  paralyzed.  Presently  he  recognized  his 
master  and  within  an  hour  from  the  time  he  took  his  distasteful  nip  he  was 
apparently  quite  normal  again." 

E.  H.  Taylor  and  H.  M.  Smith  recently  expressed  the  following  opinion: 
"We  are  convinced  that  Bitfo  marinus  Linnaeus,  as  generally  accepted, 
comprises  species  or/and  subspecies  and  is  in  fact  of  almost  generic  signifi- 
cance. However,  certain  difficulties  are  involved  in  properly  delineating  and 
naming  these  forms.  In  the  first  place  the  type  localities  of  Bufo  marines  and 
B.  aqua  are  unknown  except  that  they  are  from  the  Western  Hemisphere. 
Bufo  maculiventris  and  B.  lazarus  of  Spix  are  Brazilian,  but  lack  exact  locali- 
ties; B.  ictericus  Spix,  however,  is  cited  with  Rio  de  Janeiro  as  type  locality. 
B.  htimeralis  Daudin  'existe  dans  diverses  countries  meridional  du  nouveau 


192  HANDBOOK  OF  FROGS  AND  TOADS 

continent.'  He  mentions  one  in  the  Musee  d'Histoire  naturelle  from  Cayenne 
(French  Guiana).  This  may  be  presumed  to  be  the  type  locality  for  this 
species. 

"A  second  difficulty  is  that  seldom  are  good  series  of  these  great  toads  col- 
lected; and  the  age,  sex,  and  environmental  variations  are  known  for  only  a 
few  localities. 

"Wiegmann  described  Bufo  horribilis  from  a  series  of  cotypes  from  the 
State  of  Vera  Cruz,  and  we  are  reviving  this  designation  for  most  of  the  toads 
of  this  group  in  Mexico,  aware  that  there  are  probably  variant  populations 
even  here,  that  may  warrant  subspecific  designations'1  (Gen.,  1945,  pp.  551- 
552). 

"The  third  series  of  eggs  was  at  the  yolk  plug  stage  when  placed  in  15  and 
10%  sea  water  and  in  tap  water.  Mortality  appeared  to  be  slightly  less  in  the 
dilute  sea  water,  and  development  in  the  early  stages,  at  least,  somewhat  ac- 
celerated. 

"Twenty-seven  days  after  laying,  the  first  completely  metamorphosed  ani- 
mal was  taken  from  the  15%  sea  water  and  2  days  later  metamorphosis  was 
occurring  in  the  other  solutions.  Thereafter,  over  a  period  of  9  days,  meta- 
morphosed animals  were  taken  from  all  solutions. 

"In  general,  these  preliminary  experiments  seem  to  show  that  low  concen- 
trations of  sea  water  constitute  a  favorable  environment  for  the  development 
of  Bufo  marintis  larvae"  (C.  A.  Ely,  Gen.,  1945,  p.  256). 

"In  the  latter  part  of  November,  1939,  the  senior  author  secured  several 
specimens  of  Bufo  marinus  under  boulders  along  the  river  at  Culiacan  in 
Sinaloa.  This  was  during  the  dry  season  and  apparently  the  toads  were  hiber- 
nating. While  these  specimens  were  being  collected  a  native  boy  watched  with 
interest  as  the  toads  were  placed  in  the  sack  and  finally  commented,  'Echnn 
leche'  (they  throw  milk),  in  reference  to  the  whitish  venom  secreted  by  the 
parotoid  glands.  This  secretion  is  readily  ejected  a  distance  of  a  foot  or  so  if  the 
glands  are  squeezed.  However,  the  toads  have  not  been  observed  to  expel  the 
poison  such  a  distance  voluntarily. 

"It  has  been  noted  on  several  occasions  that  B.  marmits,  when  killed  by 
being  placed  in  formalin  or  alcohol,  commonly  secretes  quantities  of  viscous^, 
whitish  liquid  from  the  parotoid  glands"  (C.  M.  Bogert  and  J.  A.  Oliver, 
Gen.,  1945,  pp.  340-^41). 

Spotted  Toad,  Belding's  Toad,  Canyon  Toad,  Red-spotted  Toad 

Bufo  punctattis  Baircl  and  Girard.  Plate  XXXIX;  Map  13. 

Range:  South-central  Texas  west  to  Nevada,  southern  California,  Sonora, 
and  Lower  California. 

Habitat:  Desert  canyons,  breeding  in  rock-bottomed  pools  of  intermittent 
streams. 


TOADS:  BUFONIDAE 


193 


Plate  XXXVIll.  Bufo  marinus.  1,2.  Male  Plate  XXXIX.  Bufo  functatus.  i.  Male 

(X%).  3,4-  Male  (X%).  croaking  (X%)-  2,3,5-  Males  (X%).  4- 

Female  (X%). 


I94  HANDBOOK  OF  FROGS  AND  TOADS 

"Lower  California,  Aguaito  Springs,  15  miles  E.  Rosario,  1300  ft.  lat.  30°4'; 
37244;  June  9.  2  miles  NNW  Catavina,  1950  feet,  lat.  29°47/;  37245-37248; 
June  ii. 

"The  red-spotted  toad  from  Aguaito  Springs  was  the  only  individual  of 
this  species  seen  at  that  locality — a  series  of  spring-fed  pools  in  an  arid  terrain. 
Of  the  four  from  near  Catavina,  two  were  hopping  at  night  on  the  dry,  sandy 
floor  of  an  arroyo  at  least  100  feet  from  water,  and  two  were  squatting  in  a 
pool"  (L.  Tevis,  Jr.,  L.C.,  1944,  P-  6). 

Size:  Adults,  i  %-$  inches.  Males,  40-68  mm.  Females,  42-64  mm.,  or  even 
74  mm.  in  Lower  California. 

General  appearance:  Tins  is  a  small,  delicately  formed,  alert,  attractive  toad 
of  grayish,  greenish  tan,  ttiupe,  drab,  or  even  red  color,  with  a  flattened  body 
and  a  broad  flat  back  evenly  covered  with  scattered  tubercles  of  small  size. 
The  tubercles  on  the  back,  sides,  and  legs  may  be  reddish,  orange-cinnamon, 
or  light  vinaceous-cmnamon.  There  may  be  black  rings  or  partial  rings  at 
the  bases  of  the  tubercles.  The  under  parts  are  buff  or  white  and  may  be 
spotted  with  black  in  the  smaller  toads.  The  legs  may  be  barred  or  spotted 
with  black.  The  conspicuous  marks  of  this  toad  are  the  small,  round  paro- 
toids,  the  broad  interorbital  area,  and  the  sharp-edged,  often  pebbly,  canthus 
rostralis  that  gives  the  nostril  a  "boxed"  appearance.  The  eyelids  also  are  so  tu- 
bercular as  to  appear  pebbly.  It  is  a  fine  little  toad  of  very  neat,  compact  ap- 
pearance. It  frequently  gives  a  pleasant  birdhke  chirp  in  captivity.  We  picked 
up  one  and  turned  it  over.  Ir  lay  m  the  shallow  water  with  legs  drawn  up  as 
if  to  "possum";  we  never  saw  any  Bnfo  feign  this  lifeless  attitude  more  than 
this  individual. 

Color:  Female.  Hclotes,  Tex.,  March  1 3,  1925.  Upper  parts  light  brownish 
olive,'  bufTy  olive,  or  buffy  brown  on  parotoids  and  mterparotoid  area.  (The 
next 'day  the  upper  parts  in  general  were  drab.)  Upper  part  of  hind  legs  dull 
citrine  sharply  marked  from  light  under  parts  by  wax  yellow  or  primulme 
yellow.  This  color  or  cream  color  on  underside  of  thighs.  Tubercles  on  top, 
lateral,  and  ventral  surfaces  of  thighs  and  on  underside  of  fore  and  hind  feet 
are  orange-cinnamon  or  vinaccous-cinnamon  to  light  vmaceous-cinnamon. 
Under  parts  cartridge  buff,  the  area  ahead  of  hind  legs  olive-gray.  The  ins  has 
cream  or  pale  chalcedony  yellow  rim  around  the  pupil.  It  is  black  heavily" 
spotted  with  tilleul  buff,  vinaceous-buff,  and  light  vinaceous-fawn. 

Male.  Grand  Canyon,  Oct.  12,  1929.  Back  buffy  citrine  or  olive-lake,  buffy 
olive,  citrine-drab,  or  light  grayish  olive.  Legs  fore  and  hind  olive-buff.  Over 
back  are  scattered  little  spots  of  flame  scarlet,  rufous,  or  apricot  orange.  On 
sides  are  several  spots  of  black,  some  without  and  some  with  rufous  or  apri- 
cot orange  centers.  On  back  each  spot  has  such  a  center.  Black  or  grayish 
around  snout  to  eye,  more  or  less  around  tympanum,  slightly  on  upper  eye- 
lid, and  somewhat  on  parotoid.  Forelegs  spotted  with  black  but  these  spots 


TOADS:  BUFONIDAE 


'95 


with  centers  of  ground  color.  Femur,  tibia,  tarsus,  and  outer  toe  with  bars  of 
black.  Upper  labial  margin  olive-buff.  Undersides  of  hands  and  feet  and  more 
or  less  of  fore  limb  and  hind  limb  cinnamon  or  cinnamon-buff.  Throat 
honey  yellow,  old  gold,  or  tawny-olive.  Lower  throat  with  gnaphalium  green, 
tea  green,  dark  bluish  glaucous,  or  greenish  glaucous-blue.  Lower  belly  and 
most  of  buttocks  light  brownish  drab,  army  brown,  or  deep  brownish  vina- 
ceous.  Eyelids  above  and  canthus  heavily  tubercled  with  rufous  or  apricot 
orange.  Iris  rim  broken  above  and  below,  behind  and  in  front  viridine  green; 
iris  black,  heavily  dotted  and  splashed  with  ochraceous-salmon,  ochraceous- 
buff,  or  antimony  yellow.  Under  parts  white  dotted  with  small  black  dots. 


Map  13 

Another  male  has  no  spots  underneath.  A  third  has  them  on  forward  chin 
and  rear  pectoral  region,  those  on  chin  white-centered;  a  fourth  has  them 
faint  on  pectoral  region.  One  is  light  drab,  another  cinnamon;  the  drab  one  is 
avellaneous  or  wood  brown,  uniform  on  back  except  for  reddish  tubercles. 

Specimen  collected  by  V.  Bailey,  at  the  bottom  of  the  Grand  Canyon,  Sept. 
13,  1889.  "I  took  the  following  color  description  from  one  of  the  fresh  speci- 
mens (No.  16185);  Above  'malachite-green'  densely  speckled  with  small  dots 
of  bright  vermilion;  limbs  paler,  dotted  with  vermilion  and  also  with  minute 
black  specks  which  likewise  occur  on  the  flanks;  region  surrounding  nostrils 
black;  upper  lips  and  whole  under  surface  bluish  white,  irregularly  speckled 
with  black;  posterior  part  of  belly  and  underside  of  thighs  dark  brownish  flesh 
color;  soles,  dull  orange"  (L.  Stejneger,  Ariz.,  1890,  p.  117). 

Structure:  No  cranial  crests,  or  crests  obscure;  parotoids  small  and  round; 


196  HANDBOOK  OF  FROGS  AND  TOADS 

interorbital  area  broad;  ridge  from  nostril  to  eye  sharp  and  often  pebbly,  giv- 
ing the  nostril  a  "boxed"  appearance;  finger  excrescences  of  males  not  promi- 
nent; throat  of  male  somewhat  discolored. 

Voice:  The  vocal  sac  of  the  male  is  a  round  subgular  pouch.  The  call  is 
birdlike.  It  is  a  high-pitched,  yet  very  pleasing,  trill,  lasting  while  one  counts 
to  15  or  25. 

"Cape  Region  of  Lower  California.  While  collecting  with  a  light  early  in 
the  evening  forty-six  specimens  were  taken  around  the  public  square  in  the 
little  village  of  San  Antonio.  They  were  heard  calling  late  in  the  evening. 
A  specimen  captured  was  observed  to  make  a  shrill  whistling  noise  of  four  or 
five  seconds  duration,  at  intervals  of  about  the  same  length"  (J.  R.  Slevin, 
B.C.,  1928,  pp.  101-102). 

"A  long  continued  clear  trill,  resembling  that  of  a  hearth  cricket  but  with 
more  volume"  (J.  Grinnell,  MS;  and  T.  I.  Storer,  Calif.,  1925,  p.  197). 

Breeding:  They  breed  from  April  to  September,  May  being  the  most  com- 
mon month.  The  eggs  arc  single,  with  very  sticky  jelly,  and  sometimes  the 
eggs  are  stuck  together  loosely  as  a  small  film  on  the  bottom.  The  envelope  is 
single  %  inch  (3.2-3.6  mm.),  the  black  and  white  vitellus  M>rr-%o  inch  (i.o- 
1.3  mm.).  The  eggs  hatch  in  3  days  or  less.  The  small  tadpole,  i  inch  (24-25 
mm.),  has  a  very  black  body  and  a  whitish  tail  with  many  evenly  spaced  black 
dots.  The  tooth  ridges  are  %.  After  40  to  60  days,  the  tadpoles  transform  from 
June  to  August,  at  %  inch  (9-11  mm.). 

Journal  notes:  July  19,  1917,  from  5-7  P.M.  at  Texas  Pass,  Ariz.  We  heard 
plenty  of  toads  but  could  not  find  them.  At  last  P.  A.  Munz  and  I  dug  a 
croaking  male  from  beneath  a  rock.  They  croak  in  the  crevices  and  under- 
neath rocks.  In  the  pools  are  plenty  of  tadpoles  large  and  small.  In  the  evening 
we  picked  up  a  female  B.  punctatus.  In  the  canyon  were  no  end  of  males  in 
the  water  and  along  the  banks,  males  seizing  each  other.  As  yet  there  are  few 
females. 

Aug.  2.  We  started  down  Bright  Angel  Trail  at  Grand  Canyon.  At  Indian 
Gardens,  in  a  flat  shallow  area,  water  6  inches  deep,  we  found  two  egg  com- 
plements of  B.  pnnctatus  on  the  muddy  bottom.  They  were  near  the  west 
edge  in  shallow  water  and  not  under  overhanging  willows  nor  in  the  water 
cress.  The  eggs  were  more  or  less  agglutinated  on  the  bottom  about  one  egg* 
deep,  in  a  few  places  more.  Later,  we  found  many  complements  in  all  stages 
and  took  three  or  four  transformed  stages  of  Bufo  punctatits  here. 

May  6,  1925,  Helotes,  Tex.  Where  I  found  the  female  B.  punctattis  and  two 
males,  there  are  strewn  over  the  bottom  eggs,  black  and  white.  Some  are 
single;  some  touch  in  masses  but  not  strings  or  masses  which  would  stay  as 
masses  if  above  the  bottom.  These  occupy  a  foot  square  or  i  by  i^  feet.  These 
are  nearly  hatched.  On  the  rocky  bottom  they  look  like  the  black-fly  masses 
we  get  in  swift  water  in  the  Northeast.  Water  here  is  i  inch  deep.  Nothing  but 
B.  pttnctatus  was  calling  here  last  night  or  the  night  before.  We  discovered 


TOADS:  BUFONIDAE  197 

these  at  11:30  A.M.  We  photographed  them  at  i  P.M.  Just  now  under  some 
near-by  stone  a  B.  pttnctattts  male  can  be  heard. 

June  3, 1942,  St.  George,  Utah.  Went  up  on  Red  Hill  near  Dixie  sign.  In  a 
ditch  Anna  saw  tads  and  some  eggs.  She  found  a  dead  B.  functatus  in  ditch. 
Under  a  stone  I  took  a  Bufo  punctattts. 

Authorities'  corner: 
G.  P.  Englehardt,  Ariz.,  1917,  p.  6. 
N.  N.  Dodge,  Ariz.,  1938,  p.  12. 
R.  R.  Miller,  Calif.,  1944,  p.  123. 

Oak  Toad,  Oak  Frog,  Dwarf  Toad 

Bufo  quercicus  Holbrook.  Plate  XL;  Map  14. 

Range:  North  Carolina  to  Florida  west  to  Louisiana. 

Habitat:  Abundant  in  the  pine  barrens,  seeking  shelter  in  little  burrows 
shielded  by  vegetation  or  under  boards  or  logs.  Many  breed  in  shallow  cypress 
ponds  or  in  temporary  surface  rain  pools  or  ditches. 

Size:  Adults,  :J1-iJ4  inches.  Males,  19-30  mm.  Females,  20.5-32  mm. 

General  appearance:  This  pigmy  toad  has  a  light  stripe  down  the  back  and 
four  or  five  pairs  of  unconnected  spots  along  the  middle  of  back,  from  the 
first  pair  between  the  eyes  to  the  last,  which  are  merely  two  pinpoints  just 
ahead  of  vent.  They  may  be  light  brown  or  almost  black  so  that  the  spots 
barely  show.  The  skin  is  finely  roughened  with  tubercles,  many  of  which  are 
red.  This  brightly  colored  little  fellow  looks  like  a  bit  of  velvet  or  tapestry. 
The  arms  and  legs  arc  barred  with  black.  The  vocal  sac  of  the  male  is  con- 
spicuous when  deflated,  and  is  a  triangular  apron  with  the  base  on  the  gular 
line  and  the  point  extending  backward  over  the  pectoral  region.  Under  parts 
grayish  or  buffy. 

Color:  June  8,  1921.  Adult.  Dorsal  stripe  white,  pale  orange-yellow,  maize 
yellow,  sulphur  yellow,  or  cream-buff.  The  four  or  five  pairs  of  dorsal  spots  are 
black.  Upper  parts  with  some  gull  gray,  pearl,  or  pale  olive-gray.  Stripe  from 
lower  part  of  tympanum  almost  to  groin  of  the  above  grays;  also  patch  of  the 
same  back  of  angle  of  mouth,  below  tympanum,  above  arm  insertion,  in  front 
of  femur,  and  on  back  part  of  upper  eyelid.  All  these  lighter  dorsal  portions 
are  with  burnt  sienna  tubercles,  which  are  especially  prominent  along  either 
side  of  dorsal  stripe  from  hump  backward,  on  oblique  lateral  stripe,  and  on 
posterior  part  of  eyelid.  The  tubercles  on  black  areas  look  black  but  many  are 
really  burnt  sienna.  Parotoid  with  fine  and  thickly  studded  burnt  sienna  tu- 
bercles. Tubercles  on  palmar  and  solar  surfaces,  posterior  surface  of  thighs 
(partially),  groin  (a  little),  pectoral  region  (a  few  tubercles)  vinaceous- 
rufous,  Hay's  russet,  or  mars  orange.  Ventral  parts  smoke  gray,  grayish  white, 
pale  olive-buff,  or  cream  color.  Each  tubercle  stands  out:  on  venter,  close  to- 
gether and  black  between;  on  sides  and  underside  of  limbs,  wider  apart  and 


198  HANDBOOK  OF  FROGS  AND  TOADS 

intervening  black  more  apparent.  Iris  cream  color;  rim  around  pupil  and  eye 
naphthalene  yellow;  rest  largely  black  with  cartridge  buff  or  ivory  yellow. 

Structure:  Head  to  angle  of  mouth  short;  snout  pointed;  body  short;  flat; 
hind  limbs  shorter  than  body  length;  first  finger  less  than  or  equal  to  second; 
cranial  crests  divergent,  ends  connected  by  transverse  series  of  raised  warts, 
giving  the  cranial  hollow  a  parapet  behind;  parotoids  finely  spmose;  excres- 
cences on  fingers  of  male  not  prominent;  interorbital  region  broad. 

Voice:  The  vocal  sac  of  the  male  is  an  oblong  "sausage."  When  deflated  it 
is  made  up  of  folds  ot  skin  on  the  lower  throat  covered  by  a  conspicuous  apron 
or  lappet.  The  call  is  very  birdhke,  not  f  roghke.  It  is  a  very  high-pitched  whis- 
tle. The  chorus  is  deafening  and  can  be  heard  %  mile  or  more. 

May  16,  1921.  The  oak  toad  male  was  calling  before  we  approached.  He 
piped  only  low.  After  we  had  worked  him  around  for  a  photo,  he  suddenly, 
to  our  surprise,  backed  into  a  hole  at  the  base  of  a  saw  palmetto.  The  hole  was 
%  of  an  inch  in  diameter  and  not  deep.  The  note  of  this  toad  is  birdhke.  One 
will  hear  three  or  four  calls  sounding  like  those  of  piping  chickens.  Sometimes 
the  note  is  repeated  three  or  four  times.  Then  this  process  is  repeated  after  a 
short  interval.  Once  us  note  was  likened  to  that  of  a  swallow-tailed  kite.  Truly 
the  most  unfroghke  note  1  ever  heard.  It  sometimes  sounds  like  some  animal 
in  distress.  There  are  several  calling  in  the  pmey  woods.  One  calling  from 
a  tangle  of  chokebernes  (Aronta),  Qsmttnda  cinnamotnea,  Bamboo  brier 
(Smilax),  and  sweet  bays.  Couldn't  find  it.  Are  the  toads  moving  pondward? 
Later  in  the  evening  we  heard  none. 

1922.  A  combined  chorus  of  Hyla  gratiosa,  Hyla  femoralts,  Hyla  sqtttrclla, 
Chorophiltts,  Acns,  and  Bitfo  qucrcicits  left  our  ears  ringing  for  a  long  time 
after  we  left  the  pond.  By  half  closing  our  ears,  we  shut  out  Hyla  jemoralts 
and  heard  the  others  more  distinctly.  The  whole  was  a  terrible  din. 

In  1912  and  1921  some  of  the  residents  who  were  almost  invariably  accurate 
assured  me  that  the  black  snake  had  a  whistle  and  that  this  note  of  the  oak 
toad  was  the  call  in  question.  The  timed  calls  ranged  from  $2  calls  in  16  seconds 
to  10  calls  in  26  seconds. 

In  inflation  of  the  throat  these  toads  are  like  Kt/fo  comfactilu  and  Bujo  cog- 
natus.  The  lower  throat  is  the  principal  part  involved  in  the  process;  it  is 
thrown  out  into  an  elliptical  bag  or  sausagelike  balloon.  One  can  tell  when 
a  toad  is  going  to  trill  after  a  rest  because  the  body  will  inflate  to  a  large  size 
and  then  the  ludicrous  sac  projects  in  the  throat  region.  The  tip  of  the  sac 
when  not  really  inflated  comes  close  to  the  tip  of  the  chin.  Otherwise  it  ap- 
pears as  a  little,  loose,  vibrating  sac  %  cm.  out  from  the  lower  part  of  the 
throat.  When  the  sac  is  deflated,  the  body  inflates.  When  the  body  is  com- 
pressed or  deflated,  the  sac  inflates. 

Breeding:  They  breed  from  April  i  to  September  5.  It  takes  a  heavy  warm 
rain  to  start  these  little  toads  calling  vigorously.  The  eggs  are  in  bars  of  2,  3,  4, 
5,  or  6,  and  the  bars  are  from  Vrz-V*  inch  (2-7  mm.)  long  and  %o  inch  (1.3 


TOADS:  BUFONIDAE  199 

mm.)  wide.  The  eggs,  black  and  white,  }£,-•  inch  (i  mm.)  in  diameter,  are  laid 
on  the  bottom  of  shallow  pools.  The  small  tadpole  is  grayish  with  six  or  seven 
black  saddles  on  the  musculature,  and  with  heavily  marked  upper  tail  crest, 
and  the  venter  is  one  mass  of  pale  purplish  vinaceous.  They  transform  July  13 
to  August  1 6,  at  ^d-'Ke  inch  (7-8  mm.). 

Journal  notes:  May  26,  1921,  Billy's  Island,  Okefinokee  Swamp.  In  pipe- 
wort,  sedge,  and  grassy  places  at  10  A.M.  found  a  female  Eu]o  qtiercicus.  We 
hear  males  in  the  woods.  The  calls  are  more  lively  and  insistent.  Is  B.  querci- 
ciis  going  to  the  ponds  soon?  We  have  taken  four  or  five  toads  this  morning. 
Females  are  about  more  since  last  night's  thunderstorm.  In  a  burnt-over  area 
it  seemed  as  if  more  were  present.  Possibly  they  are  easier  to  find  in  this  area. 
We  found  three  males  and  three  females.  The  males  arc  not  in  holes. 

June  4,  Billy's  Island,  Okefinokee  Swamp.  About  two  inches  of  rain 
dropped,  and  the  island  seemed  teeming  with  oak  toads.  They  bred  almost 
everywhere.  All  about  the  cleared  fields,  in  piney  woods,  in  hammocks,  and 
in  numerous  other  places,  we  found  oak  toads  that  day.  On  July  3  the  species 
was  abroad  in  great  numbers.  Every  transient  shallow  pool  filled  by  the  rain 
had  them  calling.  We  took  three  or  four  pairs  and  30  to  40  males  in  short 
order. 

On  July  27, 1922,  in  a  shallow  pond,  we  heard  so  many  oak  toads  we  looked 
for  eggs.  We  found  single  bars  of  two  to  six  or  eight  eggs  rarely  attached  to 
sticks  at  the  surface,  usually  attached  to  grass  blades  0.5  to  i  or  2  inches  below 
the  surface  of  the  water,  the  water  i  to  3  inches  deep.  Other  bars  were  attached 
to  pine  needles.  Once  in  a  while  two  bars  extended  out  from  a  common  focus. 
Normally  they  were  close  together. 

Authorities'  corner: 

J.  E.  Holbrook,  Gen.,  1842,  p.  14.  P.  Viosca,  Jr.,  La.,  19233,  p.  37. 

E.  Loennbcrg,  Fla.,  1895,  p.  338.  A.  F.  Carr,  Fla.,  i94ob,  p.  54. 

H.  W.  Fowler,  Fla.,  1906,  p.  109. 

Southern  Toad,  Carolina  Toad,  Gray  Toad,  Land-Frog  (Bartram),  Land- 
Toad  (Catesby),  Latreille's  Toad,  Charming  Toad,  Hop  Toad 

Bujo  terrestns  (Bonnaterre).  Plate  XLI;  Map  9. 

Range:  North  Carolina  to  Florida,  west  to  the  Mississippi  River.  We  rather 
incline  to  the  limitation  of  this  form  to  the  coast  below  the  fall  line.  Are  the 
isolated  records  (one,  western  South  Carolina;  one,  northern  Georgia;  one, 
northern  Mississippi;  one,  northern  Louisiana;  several,  extreme  western 
Louisiana)  misidentifications  or  intermediates?  S.  N.  Rhoads  in  southeastern 
and  middle-central  Tennessee  calls  his  specimens  intermediate. 

Habitat:  Abundant  throughout  its  range,  particularly  common  in  culti- 
vated fields.  It  occurs  throughout  the  pine  barrens  and  hammocks,  in  fact  in 
any  land  habitat.  When  breeding,  it  is  usually  in  shallow  water  from  the  tiny 


166 


HANDBOOK  OF  FROGS  AND  TOADS 


Plate  XL.  Bujo  quercicus.  1,2.  Males 
croaking  (X%)-  3.8-  Males  (XO-  4,6.7.9- 
Females  (XO-  5-  Egg  bars 


Plate  XU.  Bufo  terrestrit.  i.  Mai 
(X%)-  2.  Male  (X%)-  M-  Mal«  croak 
ing  (X%).  5-  Female  (X%). 


TOADS:  BUFONIDAE  201 

pool  to  the  edges  of  lakes.  Frequently  these  toads  are  in  pools  so  transient  that 
they  can  last  but  a  few  hours,  as  in  the  furrows  in  fields  or  in  temporarily  over- 
flowed grassy  areas.  On  one  occasion  their  eggs  were  so  plentiful  in  imperma- 
nent pools  that  we  wrote  (June  4)  :  What  a  frightful  waste  of  frog  life  in  tran- 
sient pools! 

Size:  Adults,  1^-3%  inches.  Males,  42-82  mm.  Females,  44-92  mm. 

General  appearance:  These  toads  vary  in  color  from  red  or  gray  to  black. 
The  crests  on  the  head  are  prominently  raised  at  the  rear  into  clublike  promi- 
nences or  knobs.  The  skin  between  the  larger  warts  is  finely  and  uniformly 
roughened  with  tubercles  all  over,  including  the  eyelids  and  parotoids. 

Color:  June  9,  1921,  Okefinokee  Swamp,  Ga.  Male.  Line  from  front  line  of 
eyes  almost  to  vent  pale  gull  gray  or  mineral  gray.  Black  spots  along  dorsum: 
one  pair  near  cephalic  edge  of  eye;  one  pair  from  upper  eyelid  connected 
across  the  meson;  one  in  the  middorsum  between  tympana;  a  pair  either  side 
of  middle  between  the  rear  ends  of  tympana;  a  pair  of  small  spots;  a  pair  of 
large  spots  where  hump  comes;  a  pair  of  small  spots;  a  pair  just  ahead  of  vent 
— all  black  spots  of  dorsum  thinly  encircled  with  chalcedony  yellow.  More 
or  less  broken  pale  gull  gray  or  mineral  gray  line  from  tympanum  to  groin; 
below  this  a  prominent  black  area.  Tubercles  of  back  black  or  deep  brown 
tipped.  Lighter  areas  on  rear  of  hind  legs  sulphur  yellow.  Either  side  of  vent 
a  few  orange  tubercles.  Under  parts  pale  smoke  gray.  Pectoral  region,  under 
parts  of  hind  legs,  and  sides  with  black  spots.  Lower  jaw  rim  like  belly  color. 
Throat  deep  mouse  gray  or  dark  mouse  gray  with  widely  spaced  white  dots, 
giving  throat  discolored  appearance.  Light  area  of  dorsum  of  hind  foot  same 
color  as  the  rear  of  thighs.  Very  little  rusty  on  front  of  thighs  and  groin.  Top 
of  first  two  fingers  with  excrescences  and  slight  line  of  such  on  edge  of  third 
finger.  Color  of  excrescences  chocolate,  or  better,  Hay's  maroon  or  maroon. 

Female,  Lighter,  larger.  Practically  no  spots  on  pectoral  region.  None  on 
throat.  Throat  same  color  as  belly.  Practically  no  rusty  spots.  Ground  color  of 
the  dorsum  more  greenish  olive.  Sometimes  at  breeding  season  males  and 
females  may  be  alike  in  color,  e.g.,  on  April  24,  1921,  several  were  thus,  several 
pairs  reddish,  one  pair  gray.  Most  of  the  pairs,  however,  were  diverse. 

Structure:  Prominent  knobhkc  crests;  backs  of  thumb  and  second  finger 
of  male  with  excrescences,  which  are  also  on  inner  edge  of  the  third  finger; 
female  throat  usually  light;  male  throat  usually  dark;  pectoral  region  may  be 
heavily  spotted,  sometimes  only  a  median  spot. 

Voice:  The  note  sounds  much  like  our  droning  Bufo  americantts  of  the 
North,  not  like  the  scream  of  Bufo  w.  jowleri.  The  trill  is  perhaps  shorter  than 
that  of  B.  amcricanus.  The  trill  lasts  7-9  seconds,  with  intervals  of  4-60  sec- 
onds. It  is  musical  in  character.  It  has  been  described  as  a  high  trill,  a  drone, 
or  even  a  bass  roar,  for  when  many  are  calling  close  to  the  observer,  the  sound 
is  deafening.  The  choruses  can  be  heard  some  distance  away.  Like  other 
species,  they  may  give  weak  notes;  individuals  may  be  freakish,  hesitant, 


202  HANDBOOK  OF  FROGS  AND  TOADS 

shrill,  or  even,  rarely,  open  the  mouth  to  scream,  or  with  half-inflated  throat 
give  puzzling  notes.  Usually  when  croaking,  the  throat  is  distended  to  its  full 
capacity  with  the  body  compressed.  Then  the  body  is  distended  and  the  throat 
collapsed. 

The  calling  toads  in  cypress  ponds  and  bays  may  be  perched  on  a  log,  on  a 
cypress  knee,  stub,  or  stump,  at  the  base  of  a  cypress  or  gum  tree,  on  the  moss, 
or  resting  on  aquatic  plant  stems,  leaves,  or  dead  twigs,  usually  in  shallow 
water  or  at  the  edge  of  a  pond.  In  overflow  pools  they  may  be  anywhere  in 
shallow  places.  Rarely  if  ever  do  they  float  when  croaking.  This  toad  is  truly 
an  alert,  pert  animal. 

Breeding:  These  toads  breed  from  March  i,  or  earlier,  to  September.  The 
eggs  are  in  long  coils  of  jelly,  the  egg  !4v-Vi<i  inch  (1-1.4  mm.),  the  outer 
tube  Mo-%6  inch  (2.6-4.6  mm.),  the  inner  tube  Mu-%  inch  (2.2-3.4  mm-)- 
The  eggs,  separated  in  the  tube  and  with  no  partition  apparent  between 
them,  number  2500-3000,  and  hatch  in  2-4  days.  In  the  small  tadpole,  i  inch 
(26  mm.),  the  body  is  broader  toward  the  rear,  the  tail  crests  are  narrow,  the 
tail  is  short  and  rounded,  and  the  eyes  are  dorsal,  close  together;  it  is  black  in 
color.  The  tooth  ridges  are  %.  After  30-55  days,  the  tadpoles  transform  from 
April  to  October,  at  /4-%  inch  (6.5-1 1  mm.). 

Journal  notes:  April  24,  1921,  Okefinokee  Swamp,  Ga.  At  9:15  A.M.  Bitfo 
terrestris  were  calling.  Water  66°.  Sun  very  bright.  There  were  15  pairs  in  an 
area  6  feet  square.  One  pair,  both  male  and  female  reddish.  Male  about  % 
size  of  female.  Male  embraces  female  in  axillary  fashion,  the  last  two  fingers 
not  dug  into  axil  but  resting  on  arm  insertion  of  the  female.  When  the  pair  are 
not  laying,  the  male  has  the  hind  legs  free  and  floating,  but  when  the  female 
ovulates  several  inches  of  egg  string,  the  male  brings  his  knees  into  the  groin 
of  the  female  and  heels  almost  touch,  the  upper  surface  of  hind  feet  against 
the  underside  of  femur  and  near  cloacal  opening.  Female  with  hind  legs 
stretched  back  sometimes  heels  touching,  sometimes  not.  The  eggs  rest  in  the 
cup  made  by  heels  and  feet  of  male.  The  pair  may  remain  in  emission  atti- 
tude 4  or  5  minutes  or  less.  Then  the  female  crawls  i  foot  or  more.  A  minute 
may  elapse  before  another  emission.  By  11:30  A.M.  surface  of  water  was  81°. 

May  2.  One  of  the  boys  called  this  toad  "charming  toad,"  because  "it  charms 
you,  turns  your  eyes  right  green." 

June  8, 1930, 2  miles  north  of  Mandeville,  La.  In  the  pine-barren  parishes  of 
eastern  Louisiana,  Viosca  finds  Bttjo  terrestns;  the  first  one  I  espied  in  a  mixed 
magnolia  thicket  was  as  intense  a  reddish  brown  as  I  have  ever  seen  in  a  toad. 

Authorities1  corner: 
J.  E.  Holbrook,  Gen.,  1842, 5, 9. 
R.  F.  Deckert,  Fla.,  1914,  p.  2. 


TOADS:  BUFONIDAE  203 

Nebulous  Toad,  Wiegmann's  Toad,  Mexican  Toad 

Bufo  valhceps  Wiegmann.  Plate  XLII;  Map  13. 

Range:  Louisiana,  eastern  and  south-central  Texas  to  Mexico  and  Costa 
Rica. 

Habitat:  Lowlands  in  the  West  Gulf  Coastal  Plain  of  Louisiana,  in  the  hills 
in  northwestern  Louisiana,  in  the  pine  lands  of  eastern  Texas,  and  in  open 
stretches  of  streams  in  central  and  southern  Texas.  Frequently  found  in  rail- 
road ditches  or  roadside  pools. 

Size:  Adults,  2^-5  inches.  Males,  53-98  mm.  Females,  54-125  mm. 

General  appearance:  This  is  a  large  brown  toad,  with  a  light  streak  down 
the  middle  of  back  from  snout  to  vent,  a  light  area  over  each  parotoid  extend- 
ing diagonally  backward  to  the  groin.  This  light  lateral  area  is  bordered  below 
with  a  fringe  of  white  conical  tubercles.  The  skin  of  back  and  venter  is  closely 
set  with  tubercles.  There  is  a  light  line  on  the  upper  lip,  continued  beyond  the 
rear  of  the  angle  of  mouth  by  another  row  of  light  tubercles.  The  throat  of 
the  male  is  discolored — a  citrine  drab  or  water  green. 

Color:  Male.  Beeville,  Tex.,  March  12, 1925.  Parotoid  and  interparotoid  area 
wood  brown,  fawn  color,  or  cinnamon,  changing  to  drab  or  buffy  olive  on  m- 
terorbital  and  upper  eyelid,  face,  and  nasal  area.  Stripe  down  middle  of  back 
deep  olive-buff,  vmaceous-buff,  or  pale  pinkish  cinnamon.  Stripes  on  either 
side  the  same  color.  Stripe  on  upper  jaw  tilleul  buff  or  cartridge  buff.  Light 
spots  on  lower  side  the  same  color.  Belly  cartridge  buff  becoming  from  pec- 
toral region  rearward  cream-buff,  spotted  with  black.  Dark  bands  on  side  and 
back  brownish  olive.  The  dark  lateral  band  may  be  clove  brown  on  side  and 
olive-brown  above.  Two  pairs  of  black  spots  on  each  side  of  dorsal  stripe. 
Hind  legs  and  forearms  banded  with  deep  olive,  the  interspaces  wood  brown 
or  avellaneous.  Front  of  chin  cream-buff.  Pectoral  region  pale  vinaceous-fawn, 
rear  chin  discolored  citrine-drab  or  water  green.  Iris  pupil  rim  ivory  yellow  or 
pale  green-yellow  with  a  warm  sepia  line  above  it;  iris  largely  tawny,  ochra- 
ceous-orange,  or  lemon  chrome;  lower  eyelid  transparent  with  spinach  or  calla 
green  rim;  rest  of  lower  eyelid  with  scattered  spots  of  this  green. 

Female.  The  female  has  dorsal  stripe  and  light  areas  pale  olive-buff,  while 
two  males  have  them  dark  olive-buff.  Top  of  head  and  parotoids  grayish  olive 
or  light  grayish  olive.  Upper  parts  olive  or  deep  olive  with  little  of  brownish 
shades  as  in  males.  Under  parts  including  throat  deep  olive-buff,  cleaner  on 
throat  than  on  abdomen,  where  it  is  rather  dirty  in  appearance. 

April  2,  1925,  Nueces  River,  Tex.  Found  a  young  B.  valhceps  under  a  log. 
It  has  about  a  dozen  scattered,  prominent,  buff,  yellow,  or  light  cadmium  spots 
on  the  back.  Very  spotted  under  parts  and  particularly  in  midpectoral  region. 

Structure:  High  projecting  crests  on  crown  of  broad  head;  these  are  canthal, 
preorbital,  supraorbital,  postorbital,  parietal,  and  orbitotympanic  ridges;  paro- 


204  HANDBOOK  OF  FROGS  AND  TOADS 

toids  rather  small,  round,  or  triangular;  snout  obtuse;  toes  %  webbed;  inter- 
orbital  space  wide;  internasal  space  narrow;  upper  eyelid  much  less  than 
interorbital  space;  male  with  a  subgular  vocal  sac  not  revealed  by  wrinkles  on 
the  throat;  body  flat;  mouth  large;  excrescences  on  two  fingers  of  male  promi- 
nent. 

Voice:  The  vocal  sac  is  a  large,  round,  subgular  pouch.  The  call  is  louder, 
harsher,  and  lower  in  pitch  than  that  of  Bttfo  americanus.  The  croak  lasts  3-4 
seconds.  Often  the  males  take  stands  several  feet  up  from  the  pond's  edge. 

March  24-26,  Beeville.  Bufo  valliceps  calling  vigorously  along  roadside 
ditches,  in  streams  and  around  tanks  near  windmills.  Found  no  females  or 
mated  pairs.  Night  of  March  25.  Bujo  valliceps  males  are  hard  to  photograph 
at  night.  After  all  is  set  up,  they  usually  move  when  the  flashlight  is  taken  off 
preparatory  to  the  gun  flash.  What  a  time  between  the  moving  toads  and  the 
frequent  cars!  This  was  a  roadside  pool. 

March  28,  San  Diego-Alice,  Tex.  By  railroad  and  creek  bridge  heard  Bujo 
valliceps.  At  first  thought  it  was  a  toad  much  like  our  B.  americanus.  It 
sounded  something  like  it  at  a  distance,  and  not  harsh  as  it  is  when  near  by. 

Breeding:  This  species  breeds  from  March  to  August.  The  eggs  are  often 
in  double  rows  in  long  strings  of  jelly  with  the  wall  of  the  inner  tube  close  to 
the  outer.  This  jelly  grows  looser  with  age,  so  that  there  may  be  a  double  row 
of  25  to  27  eggs  in  j%  inches  (3  cm.)  or  a  single  row  of  7  to  10  eggs  in  i% 
inches  (3  cm.).  The  outer  tube  is  Y&  inch  (3  mm.),  the  inner  tube  M.O  inch 
(2.6  mm.),  and  the  vitellus  %o  inch  (1.2  mm.).  The  eggs  are  purplish  black 
and  hatch  in  iVz-2  days.  The  small  blackish  tadpole  has  8-10  black  bars  with 
intervening  pale  buff  areas  on  the  dorsum  of  the  tail  musculature.  The  tooth 
ridges  are  %.  After  20-30  days,  the  tadpoles  transform,  April  to  September, 
at  Ar-^A  inch  (7.5-12  mm.).  The  season  varies  with  the  periods  of  heavy  local 
rainfall  from  March-August.  Pope  found  a  female  ready  to  spawn  as  late  as 
August  25  in  Houston,  Tex. 

May  10, 1925,  Helotes,  Tex.  Captive  pair  laid  eggs.  Egg  strings.  Some  of  the 
eggs  seem  to  be  in  double  rows  in  string;  other  parts  of  same  string  may  be 
with  single  rows.  There  are  many  single  eggs  or  eggs  without  string  effect. 

May  13,  Helotes.  Went  to  Marnock's  Second  Crossing.  Found  B.  valliceps 
egg  strings  widely  spread  out.  These  were  long  strings  in  midwater  and  buoy- 
ant in  midplane.  They  were  more  above  the  bottom  than  eggs  of  any  toad  I 
have  seen.  One  bow  of  two  strings,  3  feet  long,  was  attached  only  at  ends. 
These  eggs  covered  an  area  of  6  or  4%  feet  square.  They  seem  to  be  the  com- 
plement of  one  female. 

June  16,  1930,  San  Benito,  Tex.  Among  the  Hypopachtis  cuneits  disgorged 
from  Thamnophis  sauritus  proximus  were  several  transformed  Bujo  valliceps, 
9, 10, 10,  10,  10.5,  n,  ii,  11.5, 12  mm.  Around  edges  of  pond  were  many  more 
but  most  were  away  from  edges  as  were  some  larger  ones  (16, 19, 19  mm.). 

Journal  notes:  April  2,  1925,  Cotulla  Free  Camp  Ground.  Here  in  an  over- 


TOADS:  BUFONIDAE 


205 


Plate  XL//.  Bufo  vallicefs.  1,2,4-  Males 
(X%).  3-  Male  croaking  (X%)« 


XLII1.  Bufo   tvoodhousii  wood- 
housii  (X%)-  i.2-  Females'  34  Males' 


206  HANDBOOK  OF  FROGS  AND  TOADS 

flow  cove  of  Nueces  River  where  water  purslane  grew  were  several  male  Bufo 
valliceps.  Saw  a  female  near  the  shore  but  was  going  away  when  we  saw  her. 
Possibly  our  light  was  responsible.  Some  10  or  15  males  were  calling  vigor- 
ously. 

April  22,  San  Benito,  Tex.  Pond  in  a  mesquite  region.  On  the  east  side  of 
this  beautiful  blue  water-lily  pond  we  found  B.  valliceps  on  the  moist  earth, 
transforming.  Soon  after  transformation  the  toads  show  the  light  line  of  tu- 
bercles on  the  side  and  the  white  spot  below  the  eye.  They  do  not  show  the 
furrowed  mterorbital  at  this  small  size. 

May  6,  Helotes,  Tex.  Tonight  at  9  P.M.,  as  we  approached  the  pond,  we 
espied  a  small  head  of  Thamnophis  proxima.  A  few  moments  later  two  larger 
water  snakes  were  close  together.  They  were  after  a  near-by  croaking  toad.  A 
little  farther  on  we  heard  two  male  B.  valliceps.  Presently  we  saw  something 
rolling  over  and  over  in  the  water.  It  was  a  water  snake.  In  the  semidarkness 
I  scooped  up  the  snake  and  all.  The  snake  dropped  a  toad.  The  toad  hopped 
limply  away.  Farther  on  we  found  a  young  Natrix  rhombifera  beside  another 
pair. 

June  16-20,  1930,  Brownsville.  All  along  the  river  in  its  high  state  heard 
B.  valliceps  at  night.  It  is  the  amphibian  note  of  the  river,  as  is  S.  campi  of  the 
dooryards. 

Authorities'  corner: 
P.  H.  Pope,  Tex.,  1919,  pp.  94-95. 

Rocky  Mountain  Toad,  Woodhouse's  Toad,  American  Toad 

Bttfo  woodhousii  woodhousii  Girard.  Plate  XLIII;  Map  14. 

Range:  Southeastern  Oregon  through  Idaho,  Montana,  South  Dakota,  to 
western  Iowa  and  western  Missouri,  south  through  Kansas,  Oklahoma,  and 
Texas  to  Mexico,  west  to  Imperial  Valley  and  up  Colorado  Valley  to  south- 
east Nevada  and  through  Utah  to  Idaho. 

Habitat:  This  toad  lives  in  canyons  in  mountains  and  on  plains  along  irri- 
gating ditches.  It  is  also  found  along  rivers  and  in  swamps.  In  fact  its  habitat 
is  very  diverse,  being  any  place  where  sufficient  moisture  obtains. 

"The  habitat  of  Bitjo  woodhousii  is  by  no  means  restricted  to  mountainous 
regions,  but  includes  surroundings  as  diverse  as  the  sagebrush  flats  of  eastern 
Montana,  the  prairie  fields  among  the  chalk  cliffs  of  western  Kansas,  the 
Hudsonian  Zone  mountain  sides  of  eastern  Colorado,  the  irrigation  ditches 
that  traverse  the  mesquite  plains  of  New  Mexico,  and  the  bottomlands  along 
the  Colorado  River  near  Yuma,  Arizona.  During  May  and  June,  according 
to  locality,  adults  of  this  species  may  be  found  breeding  in  shallow  sluggish 
creeks,  in  irrigation  ditches,  or  in  freshwater  pools  in  the  canyons"  (R.  Kel- 
logg, Gen.,  1932,  p.  74). 

Size:  Adults,  2^-4%  inches.  Males,  56-99  mm.  Females,  58.5-118  mm. 


208 


HANDBOOK  OF  FROGS  AND  TOADS 


General  appearance:  This  toad  looks  much  like  our  eastern  Bufo  a.  ameri- 
canus  but  is  larger.  The  general  color  is  grayish  drab  on  the  back,  with  several 
large  brownish  warts  that  are  usually  surrounded  by  a  slightly  larger  blackish 
area.  There  is  a  narrow,  light  middorsal  stripe.  The  sides  are  marked  with 
several  black  spots.  The  crests  on  the  head  are  dark  in  color,  but  not  particu- 
larly conspicuous.  The  tubercles  and  tips  of  fingers  and  toes  are  reddish 
brown.  The  head  is  short  and  thick. 

H.  M.  Smith  (Kans.,  1934,  p.  450),  who  has  given  us  our  best  account  of 
this  species,  contrasts  this  form  with  Bufo  a.  americantts. 


a.  atnertcantis 

1.  Warts  on  body  larger,  less  nu- 
merous. 

2.  Usually  but  one  or  two  warts  to 
a  dark  color  spot. 


3.  Skin  on  median  anterior  surface 
of  tarsi  and  metatarsi  with  black- 
ish spines. 

4.  Parotoids  broad  and  closely  ap- 
proximated,   not    separated    by 
more  than  their  own  length. 

5.  Cranial  crests  never  swollen  to 
form  a  "plateau." 

6.  Snouts  of  males  in  lateral  profile 
pointed  to  some  extent. 

7.  Belly  usually  profusely  spotted. 

8.  Song  a  high  trill  of  long  duration 
— 20-30  seconds  or  so. 

9.  Eggs  laid  single  file,  enclosed  in 
a  double  tubular  membrane,  with 
a  partition  between  each  egg. 

10.  A  median  dorsal  light  line  rarely 
present;  when  present  very  irreg- 
ular. 

n.  Parotoids  usually  separated  from 
postorbital  ridge;  the  latter,  either 
directly  or  by  a  secondary  arm,  in 
contact  with  the  tympanum. 

12.  Second  subarticular  tubercle  of 
fourth  toe  frequently  divided; 
first  but  seldom  not  divided. 


w.  woodhoiisii 

1.  Warts  on  body  smaller,  more  nu- 
merous. 

2.  Some  of  dorsal  spots  including 
many  warts  (eastern  specimens), 
or  but  one  or  two  (western  speci- 
mens). 

3.  Skin  on  median  anterior  surface 
of   tarsi   and    metatarsi   without 
blackish  spines. 

4.  Parotoids  narrow,  although  not 
so  long,  and  separated  frequently 
by  more  than  their  own  length. 

5.  Cranial  crests  frequently  swollen 
forming  a  "plateau." 

6.  Snouts  of  both  males  and  females 
sharply  truncate  in  lateral  profile. 

7.  Belly  usually  immaculate  or  with 
a  single  median  dark  breast  spot. 

8.  Song  a  low  trill  of  3-4  seconds  or 
so. 

9.  No  partitions  between  eggs;  but 
a  single  tubular  membrane. 

10.  A  median  dorsal  light  line  always 
present. 

11.  Parotoids  usually  in  contact  with 
the  postorbital  ridge,  the  tympa- 
num separated  distinctly  from  the 
latter. 

12.  Second  subarticular  tubercle  ap- 
parently never  divided;  first  but 
seldom. 


TOADS:  BUFONIDAE 


209 


Color:  Female.  Ephraim,  Utah,  Aug.  19,  1929.  Back  light  grayish  olive 
becoming  on  sides  and  fore  and  hind  legs  tea  green.  Tympanum  pea  green, 
parotoid  vetiver  green  or  light  grayish  olive.  Crests  clove  brown  or  fuscous. 
Spots  on  upper  parts  are  fuscous,  becoming  on  sides  dark  ivy  green  or  dull 
greenish  black.  No  very  regular  spots  on  either  side  of  back.  Each  fuscous 
spot  on  dorsum  wart  centered,  the  wart  being  drab  or  hair  brown.  On  sides 
spotting  more  pronounced.  Forearm  with  at  least  two  oblique  crossbars.  Tibia 
with  two  or  three  indistinct  crossbars.  Under  parts  deep  olive-buff  or  cream- 
buff.  Underside  of  hind  legs  pinkish  cinnamon.  Iris  fuscous  spotted  with 
vinaceous-cinnamon  or  pinkish  cinnamon  and  sulphur  yellow  or  marguerite 
yellow;  pupil  rim  broken  behind  and  in  front,  marguerite  yellow. 

Ephraim,  Utah,  Sept.  6,  1929.  Female.  Throat  cream  color  or  pale  chalced- 
ony yellow  becoming  warm  buff  on  breast,  cinnamon-buff  on  belly,  and  clay 
color  on  lower  side  of  femur  and  lower  belly.  Underside  of  hind  legs  reed 
yellow.  Underside  of  forelegs  like  breast.  Upper  parts  mineral  gray  or  smoke 
gray  becoming  on  fore  limbs,  side  of  face,  and  hind  legs  tea  green  or  light  gray- 
ish olive  to  grayish  olive.  Spots  on  sides  most  prominent  with  grayish  olive  or 
vetiver  green  dots  in  them.  Spots  on  dorsum  small  and  warts  not  large.  Line 
down  back  faint,  white  for  front  half.  Eye  black;  pupil  rim  sulphur  yellow  or 
light  vinaceous-fawn,  upper  eye  with  prominent  band  of  vinaceous-buff.  Inner 
metatarsal  black  or  clove  brown. 

"San  Francisco  Mts.  From  fresh  specimens:  Above  pale  olive-green  with 
a  somewhat  lighter  stripe  down  the  middle  of  the  back;  tip  of  tubercles  pale 
red  surrounded  by  black  rings;  lower  surface  olive- white"  (L.  Stejneger, 
Ariz.,  1890,  p.  116). 

Structure:  Cranial  crests,  prominent,  forming  right  angle  back  of  orbit; 
longitudinal  ridges  almost  parallel;  supratympanic  or  preparotoid  ridge  ab- 
sent; parotoid  glands  long,  slender,  divergent;  interparotoid  space  more  than 
twice  the  interorbital  space;  two  metatarsal  tubercles,  one  very  large,  one  very 
small;  throat  of  male  black  from  line  of  angle  of  mouth  forward;  first  finger 
slightly  longer  than  second;  large  warts  on  back,  each  with  several  pits;  this 
toad  is  larger  than  B.  americantts. 

Voice:  The  vocal  sac  is  a  rounded  throat  pouch.  The  call  is  a  vibrating  note 
of  high  pitch,  sweet  and  musical. 

Breeding:  This  toad  breeds  from  March  to  July.  "The  eggs  .  .  .  very 
closely  resemble  those  of  jowleri.  The  inner  membrane  is  absent,  there  being 
but  a  single  tube,  about  3.5  mm.  in  diameter.  The  egg  complement  of  a  single 
female  which  laid  in  captivity  was  25,644  by  actual  count"  (H.  M.  Smith, 
Kans.,  1934,  p.  452).  They  transform  March  25  to  September  at  %-%  inch 
(10-13.5  mm.). 

On  May  19, 1942,  at  Beaver  Dam  Lodge,  Ariz.,  we  took  about  6  adults  and 
about  23  transformed  and  transforming  specimens.  These  last  varied  from 
13.5-17  mm.  in  length,  with  a  mean  and  also  average  of  15  mm. 


210  HANDBOOK  OF  FROGS  AND  TOADS 

Journal  notes:  July  24,  1925.  At  Duncan,  Ariz.,  the  engineer  at  the  electric 
light  plant  told  me  of  several  "big"  toads  at  the  plant.  We  went  over.  They 
were  all  female  Bufo  woodhousii.  He  said  they  would  hop  up  on  the  doorsill 
and  wait  for  insects  to  drop  from  the  wall  below  the  light.  Why  are  they  all 
females?  Have  the  males  gone  to  ponds  since  the  recent  rains?  All  these  fe- 
males are  ripe,  not  spent. 

May  20,  Beaver  Dam  Lodge,  Littlefield,  Ariz.  Went  down  to  the  creek. 
Everywhere  toad  tadpoles  of  one  sort.  Never  saw  so  many.  Thousands,  hi  the 
ditches  and  in  the  stream  countless  tadpoles,  large  for  toad  tadpoles.  At  first 
I  thought  I  had  Rana  or  Scaphtopus  tads.  Many  of  these  tads  were  transform- 
ing or  transformed.  They  would  take  to  the  stream,  and  the  toadlets  had  a 
hard  time  stemming  the  current.  We  would  go  along  the  bank  and  cause 
them  to  jump  in,  and  then  we  scooped  them  with  the  net.  The  tadpoles  were 
in  the  clear  drainage  ditches,  in  the  big  pond  in  the  creek.  Here  on  the  algae, 
not  quite  emergent,  the  tadpoles  assembled  in  great  numbers.  On  the  algae 
mats  that  were  solid  the  toadlets  could  leap  along  in  fine  shape.  Went  out  in 
fields  next  to  the  creek  and  caught  ten  Bufo  woodhousii.  Tonight  at  9  o'clock 
went  to  the  bridge  and  could  hear  plenty  of  Bufo  woodhousii.  It  is  a  light  trill, 
not  feeble  but  sweet,  of  the  quality  of  the  Bufo  amcricanus  call.  They  are  not 
in  chorus.  Individual  calls  not  far-carrying.  In  the  closed  bag  the  note  of 
woodhousii  is  a  friendly  chuckle. 

July  2.  Went  up  Ramsay  Canyon,  Ariz.  Went  to  Newmans.  Here  we  saw 
B.  woodhousii  leaping  along  in  the  bright  sun  of  midday.  It  was  a  scrawny 
individual. 

July  16,  below  Albuquerque.  Last  night  at  camp  a  Bufo  woodhousii  feeding 
around  our  car. 

Authorities'  corner: 

W.  P.  Taylor,  Nev.,  1912,  pp.  344-345.       G.  A.  Moore  and  C.  C.  Rigney,  Okla., 
J.  M.  Linsdale,  Kans.,   1927,  pp.  75-         1942,  p.  78. 

76.  J.  C.  Marr,  Gen.,  1944,  p.  480. 

Fowler's  Toad,  Danver's  Toad 

Bufo  woodhousii  jowlcri  (Hinckley).  Plate  XLIV;  Map  14. 

Range:  New  Hampshire  to  eastern  Texas,  eastern  Arkansas,  Missouri, 
southeastern  Iowa,  eastward  in  Michigan  through  Ohio,  West  Virginia  to 
coast.  Extensions  up  Hudson,  Delaware,  Susquehanna,  Ohio,  and  other  rivers. 
Many  such  Pennsylvania  and  New  York  records.  Often  where  B.  a.  amcri- 
canus  and  B.  w.  jowlcri  are  co-existent  B.  w.  jowlcri  is  of  the  river,  stream 
banks,  or  lake  beaches;  B.  a.  americanus  of  the  hilly  or  mountainous  regions 
near  by.  Except  for  the  Northeast  our  range  closely  accords  with  Blair's  range. 
Intermediates  or  misidentifications  with  B.  a.  americanus,  B.  terrestris,  and 
B.  w.  woodhousii  make  its  range  difficult  to  determine  at  this  date. 


TOADS:  BUFONIDAE  211 

Habitat:  Beaches,  coasts,  lake  shores,  or  river  banks,  which  are  the  more 
sandy  and  warmer  places  throughout  its  range,  are  the  usual  choice.  It  is 
common  along  roadsides,  about  homes,  in  fields,  pastures,  and  gardens,  in 
sand  dunes  and  pine  barrens.  It  breeds  in  the  shallow  water  of  permanent 
ponds,  in  flooded  low  ground  or  roadside  ditches,  or  along  river  shores. 

Size:  Adults,  2-3/4  inches.  Males,  51-74.5  mm.  Females,  56-82  mm. 

General  appearance:  These  toads  generally  have  a  greenish  cast  with  a  yel- 
lowish or  buff  middorsal  stripe.  The  back  is  marked  by  distinct  black-edged 
dark  spots.  In  a  pair,  the  male  Bufo  jowleri  is  usually  darker  than  the  female. 
Live  toads  sent  us  from  Virginia  we  described  as  follows:  These  are  small 
toads  with  low  crests.  One  is  dull  greenish  in  color  and  one  reddish  brown, 
each  with  a  light  middorsal  line.  The  warts  are  small  and  rounded,  with  sev- 
eral grouped  in  each  dark  spot  of  the  dorsal  pattern.  The  parotoids  are  elliptic 
and  nearest  together  at  their  midpoints.  The  ventral  surfaces  are  buffy.  The 
throats  of  the  male  are  greenish  black.  One  toad  has  the  black  pectoral  spot, 
another  one  lacks  it.  Both  toads  have  dark  bands  along  the  sides.  One  has 
much  yellow  in  groin,  on  rear  of  the  femur,  and  on  tibia  and  tarsus. 

Color:  Lakewood,  N.J.,  from  W.  H.  Caulwell,  May  9,  1930.  Male.  General 
appearance  "dark  greenish."  Dorsal  color  citrine  drab,  parotoids  buffy  brown, 
drab,  or  wood  brown.  Top  of  hind  legs  and  forelegs  like  dorsum  or  deep 
olive.  Rear  of  femur,  lower  half,  dark  olive  or  black.  Rear  of  unexposed  femur 
with  pinard  yellow  or  straw  yellow.  Spots  larger  and  most  conspicuous  on 
rear  edge  of  tibia  between  bars.  Groin  with  some  barium  yellow  or  deeper 
yellow  spots.  Oblique  bar  on  upper  eyelids  meeting  or  not  on  meson.  Spot 
just  inside  of  forward  end  of  parotoid,  pair  of  spots  midway  near  meson, 
larger  pair  near  rear  end.  Last  two  pairs  sometimes  united  on  either  side.  Pair 
of  big  spots  near  meson  in  middle  of  back.  Other  pairs  frequently  present. 
These  spots,  two  bars  on  tibia  and  one  on  femur,  and  bars  on  arm  are  olive  or 
dark  olive.  Several  warts  to  each  spot  drab  or  light  grayish  olive.  Dark  spot 
on  lower  rear  of  parotoid  connected  or  not  with  irregular  lateral  band  of  dark 
grayish  olive  or  olive.  Oblique  dark  spot  from  over  tympanum  to  arm  inser- 
tion. Side  spotted  or  with  dark  lateral  band  mentioned  above.  Area  above 
band  and  dorsal  streaks  cartridge  buff,  ecru-olive,  olive-buff,  cream-buff,  or 
picric  yellow.  Eye  to  angle  of  jaw  spot  like  dorsal  ones.  Under  parts  white. 
Pectoral  spot  of  olive  or  black.  Throat  deep  grayish  olive  or  dark  grayish  olive. 
Iris  rim  reed  yellow  or  straw  yellow;  iris  black  spotted  with  light  ochraceous- 
salmon,  or  rufous. 

Female.  Dorsal  color  light  grayish  olive.  Parotoids  drab.  Bars  on  eyelids 
and  dorsal  spots  olivaceous  black  (i)  with  warts  fawn  color,  army  brown, 
or  orange-cinnamon  ("reddish").  Dorsal  streak  cartridge  buff  or  pale  pinkish 
buff.  Light  area  along  side  cream-buff  or  cartridge  buff.  Warts  on  hind  legs 
reddish  in  this  individual.  Belly  cream-buff.  Black  pectoral  spot.  Throat  like 
rest  of  under  parts. 


212  HANDBOOK  OF  FROGS  AND  TOADS 

Structure:  Crests  variable,  at  times  forming  a  boss;  adults  never  reaching 
the  greater  size  of  B.  a.  americanus;  warts  on  back  small  and  uniform;  no  pre- 
parotoid  ridge  from  parotoid  to  postorbital. 

Voice:  "The  usual  note  of  Fowler's  toad  is  a  brief,  penetrating,  droning 
scream.  Only  once  have  I  heard  a  decided  departure  from  this.  I  heard  this 
note  late  in  April  in  Gwinnett  Co.,  in  upper  Georgia.  A  single  individual  of 
a  noisy  congregation  of  males  had  the  unmistakable  trill  of  the  common  toad, 
but  short  and  decisive  like  the  Fowler's  song.  It  was  a  perfect  combination  of 
the  notes  of  both"  (H.  A.  Allard,  Ga.,  1908,  pp.  655-656). 

"While  we  do  not  agree  with  Mr.  Allard  in  calling  the  song  of  Fowler's 
toad  a  'scream'  or  'wail/  it  certainly  has  much  less  music  to  it  than  the  trill  of 
the  American  toad.  The  notes  are  more  closely  connected,  so  that  a  sort  of 
buzzing  is  produced"  (W.  DeW.  Miller  and  J.  Chapin,  N.J.,  1910,  p. 


"A  male  toad  is  a  persistent  singer  during  its  stay  in  the  water.  Its  song  is 
a  combination  of  a  low  whistle  and  a  moan,  and  the  two  sounds  do  not  melt 
into  a  chord.  The  combined  sound  is  discordant  and  decidedly  unpleasant 
to  a  musical  ear,  but  at  a  distance  the  sound  is  more  pleasant,  for  the  moan 
is  not  apparent  and  only  the  whistle  is  heard.  The  sound  lasts  from  two  to 
three  seconds,  and  may  be  repeated  at  intervals  of  about  ten  seconds.  In  1911 
many  sang  in  the  daytime,  but  in  1912  and  1913  very  few  were  heard  except  at 
night"  (F.  Overton,  N.Y.,  1914,  p.  27). 

"One  of  the  most  striking  differences  between  the  two  species  lies  in  their 
voices.  While  that  of  the  Common  Toad  is  high-pitched  and  musical,  the  note 
of  Fowler's  Toad  is  nasal,  and  lower  in  pitch.  Like  the  voice  of  the  Common 
Toad,  it  carries  well  and  may  be  heard  at  a  considerable  distance"  (K.  P. 
Schmidt,  111.,  1929,  p.  9). 

Breeding:  This  form  breeds  from  April  15  or  earlier  to  mid-August.  In  a 
given  locality  it  breeds  later  than  the  American  toad.  The  eggs  are  in  long 
files,  crowded  at  first  in  a  double  row,  and  numbering  as  many  as  8000.  The 
egg  is  %r>-M  o  inch  (1.0-1.4  mm-)>  the  outer  tube  1/io-'/'i  «  inch  (2.6-4.6  mm.), 
the  inner  tube  absent.  The  tadpole  is  small,  \V\  2  inches  (27  mm.),  its  greatest 
width  toward  the  rear  of  the  body.  The  tail  crests  are  low,  the  tooth  ridges  %. 
After  40  to  60  days,  the  tadpoles  transform  from  mid-June  to  August  or  later 
at  %«-%  inch  (7.5-11.5  mm.). 

"The  egg  strings,  which  resemble  those  of  the  American  toad  except  that 
the  gelatinous  tube  shows  no  distinct  inner  layer  nor  partitions  separating  the 
eggs,  have  been  noted  in  Ohio  as  early  as  May  and  as  late  as  June  24"  (C.  F. 
Walker,  Ohio,  1946,  p.  35). 

Notes  by  S.  C.  Bishop  and  Walter  Schoonmaker:  "The  eggs  of  the  Fowler's 
toad,  Bujo  jowleri  (Carman),  are  laid  later  than  the  American  toad,  Bufo 
americanus  (Holbrook).  We  collected  some  of  the  eggs  of  the  Fowler's  toad 
on  the  i3th  of  May,  1924,  at  Raft's  Pond,  near  Albany,  New  York.  From  these 


TOADS:  BUFONIDAfi 

we  made  the  following  observations.  The  eggs  of  the  Fowler's  toad,  unlike 
the  American  toad,  are  sometimes  laid  in  the  double  string. 

"May  14.  The  eggs  were  segmented.  Later  they  became  elongated. 

"May  15.  Hatched.  The  tadpoles  were  hanging  on  by  their  suckerhke  mouth 
parts,  tails  down. 

"May  1 6.  Grown  considerably  larger. 

"May  17.  Length  7  mm.  Gills  present. 

"May  18, 11:30  A.M.  Actual  length  8.5  mm.  Gills  larger. 

"May  19, 9:55  A.M.  Length  10.5  mm.  Gills  gone  from  both  sides.  Mouth  parts 
developed. 

"May  20.  Length  11.5  mm, 

"May  21.  Length  12.5  mm. 

"May  26.  Length  14  mm. 

"June  12.  Length  20.5  mm.  Hind  legs  present. 

"June  16.  Length  21  mm.  Hind  legs  very  well  developed. 

"June  20.  Length  21  mm. 

"June  24.  Length  21  mm.  Hind  legs  well  developed.  Front  legs  also  devel- 
oped. Shape  of  body  changed  greatly,  also  smaller.  Tail  beginning  to  shorten. 

"June  28.  Tail  nearly  gone  m  some  specimens  and  in  others  the  tail  is  en- 
tirely gone.  Some  specimens  grow  faster  than  others.  The  young  now  look 
like  the  adult. 

"On  May  19, 1924,  at  the  pond  more  eggs  were  laid,  embryos  elongated  May 
2ist,  and  hatching  May  22nd,  and  July  1-2,  little  toads  leaving  ponds.  Males 
began  calling  May  13,  heard  May  19,  noted  calling  at  intervals  until  July  2." 

The  pair  brought  to  the  authors  were  near  the  maximum  sizes  for  the  spe- 
cies, the  male  being  70  mm.  and  the  female  8n  mm.  in  body  length. 

Journal  notes:  June  i,  1917.  About  6  miles  beyond  Dmwiddie,  Va.,  near  the 
road,  found  several  files  of  toad's  eggs.  In  one  case  the  string  was  strung  out 
in  a  file  8  or  10  feet  long  in  the  current.  In  another  case,  in  a  backwater,  the 
mass  was  tangled  around  sticks.  In  most  cases  the  file  seems  to  contain  a 
double  row  of  eggs.  The  note  we  questioned  yesterday  evening  was  Bufo 
jowleri.  It  is  quite  different  from  the  sweet  droning  note  of  Bttfo  americanus. 
Went  out  at  night  with  a  flashlight  and  captured  many  B.  jowleri  and  a  few 
mated  pairs.  One  pair  laid  in  the  fish  can  overnight.  The  embrace  is  axillary. 

April  15,  1921.  Air  temperature  68°F.  Tonight  at  9  o'clock,  Drs.  Vernon 
and  Julia  Haber  took  me  out  to  the  Oakwood  Cemetery,  East  Raleigh,  N.C., 
and  to  St.  Augustine  grounds.  Just  beyond,  we  heard  a  chorus  of  what  I  at 
first  mistook  for  the  "bleat"  of  Microhyla  carohnensis.  It  surely  is  not  the 
note  of  our  northern  Bufo  americanus,  which  is  a  sweeter  drone  than  that  of 
B.  jowleri.  In  shallow  water  along  a  little  drain  were  plenty  of  them.  The  call 
is  a  striking  nasal  whir-r-r-r.  The  Habers  say  "like  a  lamb." 

April  15,  Raleigh,  N.C.  Fowler's  toads  were  still  in  chorus  and  strongly 
breeding,  though  started  much  earlier.  The  male  of  the  mated  pair  has  the 


214 


HANDBOOK  OF  FROGS  AND  TOADS 


first  two  fingers  doubled  back  and  dug  into  the  axils  of  the  female.  Often  the 
other  two  fingers  may  not  be  doubled  back  but  lie  next  to  the  belly  of  the 
female,  or  sometimes  these  two  fingers  will  rest  on  the  shoulder  insertion  with 
only  the  first  two  in  the  axil  proper.  The  pairs  brought  in  were  not  laying  at 
3  A.M.  but  at  6:45  A.M.  they  were  well  along  in  oviposition.  Water  temperature 
at  which  they  were  laying  is  67 °F. 

April  16,  St.  Augustine  grounds,  Raleigh,  N.C.  In  the  stream  near  the  edge 
found  Fowler's  toad  eggs  wrapped  around  plants.  They  were  in  shallow  tur- 
bid areas,  cattle-punched,  and  in  water  2  to  4  inches  deep;  many  were  laid 
last  night.  They  were  among  speedwell,  chickweed,  smartweed,  marshy  St.- 
John's-wort,  and  other  plants,  hardly  any  of  which  were  more  than  3  or  4 
inches  high  at  this  season. 

June  u,  1930,  Tickfaw  River  country,  La.  At  night  found  Bufo  valltceps 
and  Bufo  jowlen  in  same  woods.  Found  and  heard  B.  fowlen  in  ditches.  This 
form  is  Bufo  jowlen  in  voice  and  structural  characters;  in  some  ways  not 
typical  in  color.  Viosca  says  that  Bufo  valltceps  and  Bufo  fowlen  may  possibly 
interbreed  in  Louisiana. 

Authorities'  corner:  Much  printer's  ink  has  been  employed  on  the  differ- 
ences between  B.  w.  jowlen  and  Bufo  a.  amertcanus.  No  one  has  known  the 
form  more  intimately  than  W.  DeWitt  Miller.  Two  authors  who  helped 
clarify  the  differential  characters  are  G.  S.  Myers  (Ind.,  1927,  pp.  50-53)  and 
M.  G.  Netting  (Gen.,  1930,  pp.  437-443).  At  the  same  time,  1927-1930,  we  in- 
dependently assembled  these  characters  in  manuscript  (15  pp.).  We  regret 
that  space  necessitates  the  elimination  of  this  material,  but  we  include  refer- 
ences on  the  points  of  discussion. 

READING  REFERENCES  TO  1930 


1.  Voice 

a.  S.  Garman,  Check  List,  1884,  p. 
42. 

b.  F.  Overton,  N.Y.,  1914,  p.  27. 

c.  M.  Brady,  Va.,  1927,  p.  27. 

d.  Almost    every    author    on    the 
species. 

2.  Crests 

a.  S.  Garman,  C.L.,  1884,  p.  42. 

b.  E.  D.  Cope,  Gen.,  1889,  pp.  278, 
279. 

c.  M.  C.  Dickerson,  Gen.,  1906,  p. 

97- 

d.  R.  F.  Deckert,  Gen.,  1917,  p.  114. 

e.  E.  R.  Dunn,  N.C.,  1917,  p.  621. 

f.  M.  G.  Netting,  Gen.,  1930,  p. 

439- 


Size 

a.  S.  Garman,  sec  above. 

b.  O.  P.  Hay,  Ind.,  1892,  p.  458. 

c.  W.  DeW.  Miller  and  J.  Chapin, 
N.J.,  1910,  p.  458. 

d.  E.  R.  Dunn,  N.C.,  1917,  p.  621.    * 

e.  M.  Brady,  Va.,  1927,  p.  27. 
Head 

a.  E.  D.  Cope,  Gen.,  1889,  p.  278. 

b.  A.  G.  Ruthven,  Mich.,  1917,  p.  4. 
Length  of  leg 

a.  O.  P.  Hay,  Ind.,  1892,  pp.  458- 

459- 

b.  R.  F.  Deckert,  Gen.,  1917,  p.  114. 

c.  A.  G.  Ruthven,  see  above. 

d.  W.  DeW.  Miller  and  J.  Chapin, 
N.J.,  1910,  p.  316. 


TOADS:  BUFONIDAE 


215 


e.  M.  G.  Netting,  Gen.,  1930,  p.  438. 

6.  Color 

a.  O.  P.  Hay,  Ind.,  1892,  p.  458. 

b.  M.  C.  Dickerson,  Gen.,  1906,  p. 
96. 

c.  W.  DeW.  Miller  and  J.  Chapin, 
N.J.,  1910,  p.  316. 

d.  R.  F.  Deckert,  Gen.,  1917,  pp. 
113,114. 

e.  E.  B.  S.  Logier,  Ont.,  1925,  p.  94. 

f.  G.  I.  Myers,  Ind.,  1927!),  pp.  51, 

5*. 

g.  K.  P.  Schmidt,  111.,  1929,  pp.  8-9. 

h.  Many  more  writers. 

7.  Vertebral  line 

a.  O.  P.  Hay,  Ind.,  1892,  p.  458. 

b.  M.  G.  Netting,  1930,  p.  441. 

8.  Dorsal  pattern 

a.  M.  C.  Dickerson,  Gen.,  1906,  p. 

96. 

b.  G.  S.  Myers,  Ind.,  1927^  pp.  51- 

52- 

9.  Under  parts 

a.  M.  C.  Dickerson,  Gen.,  1906,  p. 
96. 

b.  H.  A.  Allard,  Ga.,  1907,  p.  381. 

c.  W.  DeW.  Miller  and  J,  Chapin, 
N.J.,  1910,  p.  316. 

d.  E.  R.  Dunn,  N.C.,  1917,  P-  622. 

e.  G.  S.  Myers,  Ind.,  1927^  p.  52. 

f.  M.  G.  Netting,  Gen.,  1930,  p.  441. 
10.  Iris 

a.  W.  DeW.  Miller  and  J.  Chapin, 
N.J.,  1910,  p.  316. 

b.  A.  G.  Ruthven,  Mich.,  1917,  p.  4. 

c.  G.  S.  Myers,  Ind.,  1927^  pp.  51- 
52. 

n.  Warts 

a.  M.  C.  Dickerson,  Gen.,  1906,  p. 

97- 


Plate  XLIV.  Bufo  woodhousii  fowlerL 


b.  H.  A.  Allard,  Ga.,  1907,  p.  304. 

c.  W.  DeW.  Miller  and  J.  Chapin,      ^,5-  Males  (X%).  2.  Adult  (Xtt).  3- 
XT  T  *  remale 

N.J.,  1910,  p.  317- 

d.  E.  R.  Dunn,  N.C.,  1917,  p.  621. 

e.  R.  F.  Deckert,  Gen.,  1917,  p.  114- 

f.  G.  S.  Myers,  Ind.,  1927^  p.  51. 

g.  K.  P.  Schmidt,  111.,  1929,  p.  9. 


2l6 


HANDBOOK  OF  FROGS  AND  TOADS 


h.  M.  G.  Netting,  Gen.,  1930,  p.     17.  Eggs 


440. 
12.  Muzzle 

a.  M.  C.  Dickerson,  Gen.,  1906,  p. 


1 8.  Palettes 

a.  M.  G.  Netting,  Gen.,  1930,  pp. 


439>  44°- 
96.  19.  Range 

b.  R.  F.  Deckert,  Gen.,  1917,  p.  114.  a.  K.  P.  Schmidt,  La.,  1920,  p.  85. 

c.  E.  R.  Dunn,  N.C.,  1917,  p.  621.  b.  G.  S.  Myers,  Ind.,  1927^  pp.  51, 

52. 
c.  Many  more  writers. 


d.  M.  G.  Netting,  Gen.,  1930,  p. 

440. 
13.  Parotoids  20.  Bars  on  hind  legs 

a.  M.  C.  Dickerson,  Gen.,  1906,  pp.  a.  M.  G.  Netting,  Gen.,  1930,  p.  441. 
96,  97.                                            21.  Supra-anal  warts 

b.  A.  G.  Ruthven,  Mich.,  1917,  p.  i.  a.  M.  G.  Netting,  Gen.,  1930,  p.  441. 

c.  E.  R.  Dunn,  N.C.,  1917,  p.  621.        22.  Hybridization 

a.  W.  DcW.  Miller  and  J.  Chapin, 
N.J.,  1910,  p.  317. 

b.  C.  L.  Hubbs,  111.,  1918,^43. 

c.  A.  L.  Pickens,  S.C.,  19273,  pp. 


d.  K.  P.  Schmidt,  111.,  1929,  p.  9. 

14.  Shape 

a.  H.  A.  Allard,  Ga.,  1907,  p.  384. 

15.  Odor 

a.  W.  DeW.  Miller  and  J.  Chapin, 
N.J.,  1910,  p.  316. 

1 6.  Breeding  season 

a.  All  authors  agree  the  breeding 
season  is  2-4  weeks  later  than  in 


25,  26. 

d.  A.  L.  Pickens,  S.C.,  1927^  p. 
109. 

e.  M.  G.  Netting,  Gen.,  1930,  p. 
442. 


Bufo  a.  amcricanus. 

For  five  recent  papers  on  B.  w.  jowleri,  see  A.  P.  Blair  (Gen.,  i943a),  M.  G. 
Netting  and  C.  J.  Coin  (Fla.,  1945),  C.  F.  Walker  (Ohio,  1946),  R.  A.  Littleford 
(Md.  1946),  and  A.  P.  Blair  (N.J.,  1947)- 


FAMILY  HYLIDAE 

Genus  ACR1S  Dumeril  and  Bibron 
Map  15 

Cricket  Frog,  Savannah  Cricket,  Cricket  Hylodes,  Peeper,  Southeastern 
Cricket  Frog,  Cricket  Hyla,  Sphagnum  Cricket  Frog,  Coastal  Cricket  Frog 

Acris  gryttus  gryllus  (Le  Conte).  Plate  XLV;  Map  15. 

Range:  Dunn  (Gen.,  1939,  PP- 153~154)  g^ves  southeastern  Virginia  (Nor- 
folk) to  Florida  to  parishes  of  Louisiana. 

Habitat:  Terrestrial,  shade-loving  frog.  In  meadows  or  about  creeks  or 
ponds  in  the  open  vegetation  mats  or  wooded  edges. 

"All  in  the  region  of  coastal  sphagnum  swamps"  (C.  S.  Brimley,  N.C., 
1940,  p.  15). 

"The  most  abundant  amphibian  of  the  region  and  the  only  one  which 
could  be  taken  during  every  month  of  the  year.  Each  roadside  pool  or  ditch 
contained  numerous  individuals  and  the  swamps  and  marshes  literally 
swarmed  with  them.  In  the  short  winter  season,  here  lasting  from  about  the 
middle  of  December  till  the  first  week  or  two  in  February,  Cricket  Frogs 
were  much  less  in  evidence  than  at  other  times,  and  as  collecting  is  then  at  its 
easiest  it  is  clear  that  the  majority  hibernate.  But  even  on  the  coldest  winter 
days  no  difficulty  attends  the  capture  of  a  plentiful  supply,  which  thus  pro- 
vides a  dependable  source  of  food  for  many  of  the  snakes  kept  in  laboratory 
cages  throughout  the  year.  Sporadically  met  with  in  the  piedmont"  (J.  D. 
Corrington,  S.C.,  1929,  p.  65). 

"Almost  any  aquatic  situation;  commonest  in  marshes  and  ditches  with 
shallow  margins  choked  with  hydrophytes"  (A.  F.  Carr,  Jr.,  Fla.,  i94ob, 

P-55)- 

Size:  Adults,  %-i%  inches.  Males,  15-29  mm.  Females,  16-33  mm- 
General  appearance:  This  is  a  small  tree  frog,  but  it  looks  like  a  small  true 
frog  (Rana).  It  varies  in  color,  black,  dark  brown,  reddish  brown,  light 
brown,  green,  or  gray;  or  the  markings  may  be  reddish  on  a  green  ground. 
Between  the  eyes  there  is  usually  a  dark  triangle,  white-bordered  behind.  This 
species  possesses  rear  femoral  stripes,  oblique  bars  on  the  sides,  light  spots  on 
the  jaw,  and  an  oblique  white  stripe  from  the  eye  to  the  arm.  The  skin  is 
more  or  less  tubercular. 
"Much  like  the  preceding  [A.  g.  crepitans},  but  the  head  is  longer  and 


2l8 


HANDBOOK  OF  FROGS  AND  TOADS 


more  pointed,  the  dark  triangle  between  the  eyes  is  longer  and  more  acute 
behind;  the  edge  of  the  upper  jaw  is  dark  with  three  or  four  light  vertical  bars 
or  spots  on  each  side  and  the  dark  stripe  on  the  back  of  the  thigh  is  darker 
and  more  constant.  Slightly  smaller  and  slimmer  than  the  common  cricket 
frog.  This  species  averages  darker  and  smaller  than  the  preceding,  the  darker 
color  probably  owing  to  the  darker  soil  of  the  region  it  inhabits,  and  I  have 
seen  but  few  wholly  green  specimens.  In  habits  it  is  similar,  being  quite  as 


Map  75 

active  if  not  more  so.  Both  species  occur  in  both  shady  and  sunny  situatioas. 
By  Cope  this  was  considered  a  more  Southern  race  of  the  preceding,  but  most 
authors  have  recognized  it  as  merely  an  individual  variation.  Dr.  Percy 
Viosca,  Jr.,  however,  seems  to  have  shown  fairly  conclusively  that  they  are 
two  distinct  species  and  I  agree  with  him.  Dunn  (1939)  says  the  difference 
in  the  amount  of  the  webbing  on  the  hind  feet  is  the  best  criterion  by  which  to 
separate  the  two,  and  I  find  in  looking  over  some  fifty  specimens  in  the  State 
Museum,  about  evenly  divided  between  the  two  forms,  that  a  glance  at  the 
first  or  shortest  toe  is  usually  sufficient  to  distinguish  either;  if  the  webbing 
extends  to  the  tip,  the  frog  is  crepitans,  if  the  last  joint  is  free  from  web  it  is 
gryttus"  (C.  S.  Brimley,  N.C.,  1940,  p.  15). 


TREE  FROGS:  HYLIDAE 


219 


Color:  On  one  day,  April  23,  1921,  in  Georgia,  I  saw  black,  dark  brown, 
reddish  brown,  light  brown,  green,  and  gray  specimens  of  Acris. 

Okefinokee  Swamp,  Ga.,  June  25, 1921.  Male.  Stripe  down  back  and  around 
triangle  dark  olive-buff.  Throat  raw  sienna.  All  over  the  throat  are  collections 
of  dark  dots,  sometimes  arranged  in  a  reticulate  fashion.  Iris  pale  vinaceous- 
drab  on  black,  light  orange-yellow  pupil  rim. 

Female.  Clove  brown  above;  triangular  spot  between  eyes  obscured  by  this 
dark  color;  throat,  breast,  and  belly  pale  olive-buff;  more  or  less  of  same  color 
on  underside  of  forelegs,  the  spot  below  eye,  along  the  upper  and  lower  jaw, 
and  the  line  from  eye  to  arm  insertion;  area  back  of  arm  insertion  pallid  vma- 
ceous-drab;  oblique  bar  on  side  clove  brown  with  pale  olive-buff  and  olive- 
ocher;  underside  of  hind  legs  clear  with  little  pigment;  long  stripe  on  rear  of 
femur,  snuff  or  Dresden  brown  with  clear  unpigmented  stripe  below  and 
above;  also  another  brown  stripe  above  the  upper  clear  area;  from  vent 
around  bases  of  hind  legs  to  venter  are  pale  olive-buff  papillae. 

Structure:  Tympanum  indistinct;  tympanic  fold  present;  fold  across  breast 
frequently  present;  disk  small;  hind  limb  very  long;  tibia  very  long. 

"Width  of  head  across  the  base  of  lower  jaw  less  than  its  length  from  that 
point  to  tip  of  snout,  legs  longer  and  hind  feet  less  webbed,  the  web  of  the  first 
toe  absent  from  the  last  joint;  heel  when  leg  is  extended  reaching  beyond  the 
tip  of  the  snout.  Sphagnum  cricket  frog  (Acris  gryllus)"  (C.  S.  Brimley,  N.C., 
1940,  p.  15). 

"Webbing  on  toes  less  extensive;  last  three  phalanges  of  fourth  toe  free  from 
webbing;  usually  with  third  and  fifth  toes  not  reaching  middle  of  the  third 
phalanx  from  tip  of  fourth  toe.  Acris  g.  gryllus"  (E.  B.  Chamberlain,  S.C., 
1939,  p.  12). 

Voice:  Characterizations  of  the  Acris  call  are  numerous. 

April  1 6, 1921,  at  Raleigh,  N.C.  Acris  gryllus  call  sounds  like  a  rattle  or  some 
of  the  metal  clickers,  gtcl^t  gtcf(,  gicf(f  or  l^ic^  f(tc^f  fycl(,  in  rapid  succession. 
Frequently  one  finds  the  males  with  inflated  vocal  sacs  even  when  not  calling. 
When  calling,  the  throat  is  never  fully  deflated.  After  a  call  it  may  be  swollen 
to  three-quarters  its  full  capacity.  Then  when  the  call  is  given,  the  body  sides 
are  compressed  and  the  vocal  sac  is  extended  to  its  limit. 

Holbrook  (Gen.,  1842,  IV,  132)  wrote,  "This  is  a  merry  little  frog,  constantly 
chirping  like  a  cricket,  even  in  confinement.  .  .  ."  Of  his  captives  he  said, 
"Their  chirp,  at  times,  was  incessant  and  sprinkling  them  with  water  never 
failed  to  render  them  more  lively  and  noisy." 

"Call  an  irregular  series  of  iiks;  comparable  to  sound  made  by  scratching 
teeth  of  a  comb;  somewhat  cricket-like;  ik-ik,  ii-ik,  ii-ik"  (A.  F.  Carr,  Jr.,  Fla., 
1934,  p.  21). 

Breeding:  This  species  breeds  from  February  to  October.  The  single  eggs  are 
few  (250),  are  brownish  and  white,  and  are  attached  to  stems  of  grass  in  shal- 
low water  or  are  strewn  on  the  bottom.  The  egg,  %5  inch  (0.9-1.0  mm.),  has 


220  HANDBOOK  OF  FROGS  AND  TOADS 

a  single  envelope  Yio-Vs  inch  (2.4-3.6  mm.).  The  dark  olive  tadpole  is  me- 
dium, i1Me  inches  (42  mm.),  full  and  deep-bodied,  its  tail  long  with  acumi- 
nate tip  and  with  a  black  flagellum.  The  tooth  ridges  are  %.  After  50  to  90  days 
or  longer,  the  tadpoles  transform  from  April  to  October  at  %-%  inch  (9-15 
mm.). 

June  i,  1917,  at  Dinwiddie,  Va.,  we  found  this  species  breeding.  They  had 
chosen  a  shallow  grassy  meadow  pool  i  to  4  inches  deep.  The  eggs  were  at- 
tached singly  to  sedge  stems  or  were  strewn  singly  on  the  bottom.  In  one  or 
two  cases,  three  or  four  eggs  were  close  together.  Many  of  the  eggs  were  in 
water  not  more  than  an  inch  deep.  The  eggs  are  firm.  We  found  no  more  than 
ten  eggs. 

"Calling  was  earliest  heard  on  January  22.  This  increased  to  the  proportions 
of  a  chorus  by  March  17.  Clasping  was  first  observed  on  April  19  while  the 
first  eggs  were  procured  on  April  25"  (B.  B.  Brandt,  N.C.,  19363,  p.  217). 

"They  may  breed  during  any  month  of  the  year.  The  eggs  are  laid  in  very 
shallow  water,  often  among  semiaquatic  or  terrestrial  vegetation  which  has 
been  temporarily  inundated"  (A.  F.  Carr,  Jr.,  Fla.,  1940,  p.  55). 

Journal  notes:  The  Cornell  party  of  Dec.  22,  I9i3~jan.  i,  1914,  found  this 
species  active  and  took  several  specimens.  Doubtless  this  species  is  more  or  less 
active  throughout  the  year  in  the  Okefinokee  Swamp. 

April  23,  1921,  Okefinokee  Swamp.  Acris— captured  a  lot  of  them.  Some- 
times black  on  a  black  soil  and  hard  to  see  except  when  they  jump.  Some  brown 
all  over  back  (except  for  dark  marks)  when  on  brown  pine  needles.  Sometimes 
green  all  over  except  for  the  dark  marks.  Sometimes  gray  over  drier  sand. 
Among  some  of  the  light  brown  needles  Acris  reddish  brown  even  on  back 
of  fore  limbs  and  hind  limbs. 

A  few  of  our  notes  on  the  jumping  records  of  Acris  follow: 

April  23,  1921.  Acris  usually  jumps  for  several  leaps  before  it  disappears. 
April  25.  Acris  males  jump  3  feet  at  a  time  on  the  water's  surface.  May  17.  Acris 
can  leap  at  least  my  own  pace.  May  21.  Acris  has  been  hopping  around  on  the 
ground  and  into  small  bushes  from  the  ground  and  down  to  the  ground  again. 

Authorities'  corner: 

W.  Bartram,  Gen.,  1791,  p.  278.  C.  J.  Goin,  Fla.,  1943,  p.  148. 

P.  Fountain,  Ga.,  1901,  p.  63.  P.  Viosca,  Jr.,  La.,  1944,  p.  55. 

R.  F.  Deckert,  Fla.,  19153,  p.  22.  G.  L.  Orton,  La.,  1947,  pp.  377-378. 

A.  F.  Carr,  Jr.,  Fla.,  1940,  p.  55. 

Cricket  Frog,  Western  Cricket  Frog,  Valley  Cricket  Frog,  Western  Cricket, 
Peeper,  Savanna  Cricket  Frog,  Rattler 

Acris  gryllus  crepitans  Baird.  Plate  XL VI;  Map  15. 

Range:  Dunn  (Gen.,  1939,  pp.  153-154)  gives  Connecticut,  s< 
New  York,  New  Jersey,  Pennsylvania,  Maryland,  Delaware,  northwest  to 


TREE  FROC&:  HYLIDAE 


221 


Plate  XLV.  Acris  gryllu;  gryttus  (XO-  ***   ^LVL   Acris 

1,3.  Females.  2.  Male.  4.  Tadpole.  (X%)-  »A5A7-  ^males.  3,4-  Males.  8,9. 

Tadpoles. 


HANDBOOK  OF  FROGS  AND  TOADS 

Canada,  west  to  Utah  and  New  Mexico,  and  south  through  Virginia  to 
Georgia  and  western  Texas.  Sea  level  to  2000  feet. 

Habitat:  Swales,  lake  margins,  stream  edges,  springs,  pasture  pools. 

"This  tiny  frog  is  distributed  all  over  the  State  wherever  there  are  lakes, 
ponds,  springs,  or  streams.  I  have  found  it  even  in  the  heart  of  well  populated 
cities  in  little  pools  formed  by  rains.  While  allied  to  the  true  tree  frogs,  this 
species  never  climbs  trees  but  lives  among  water  plants  and  in  the  vegetation 
along  shore.  When  alarmed  it  retreats  to  the  water  after  the  manner  of  a  true 
water  frog"  (J.  K.  Strecker,  Jr.,  Tex.,  1915,  p.  49). 

"This  is  one  of  the  commonest  forms  found  in  the  Wichita  National  Forest 
and  if  an  effort  were  made  large  numbers  could  be  taken.  Several  dozen  were 
preserved  on  this  trip  and  some  hundreds  are  in  the  collections  as  the  result  of 
many  short  collecting  trips  which  have  been  taken  to  this  forest.  They  are 
found  almost  anywhere  that  water  was  available  and  in  this  region  were  plenti- 
ful along  most  parts  of  the  streams  and  particularly  around  the  edge  of  Lost 
Lake  and  other  slow-flowing  water  areas.  Their  abundance  along  West  Cache 
Creek  in  spite  of  numerous  campers  was  very  noticeable"  (A.  I.  Ortenburger 
and  B.  Freeman,  Okla.,  1930,  p.  177). 

"It  is  generally  abundant  wherever  permanent  waters  are  to  be  found,  where 
it  may  be  taken  as  early  as  the  middle  of  February  and  as  late  as  November" 
(D.  A.  Boyer  and  A.  A.  Heinze,  Mo.,  1934,  p.  189). 

Viosca  stated  that  Acns  gryllus  abounded  along  the  creeks  of  his  Pine  and 
Hardwood  Uplands  division  or  "Shortleaf  Pine  Hills."  "Fresh  Water  Marshes: 
Acris  crepitans,  although  a  lowlands  species  generally,  is  especially  abundant 
here  .  .  ."  (P.  Viosca,  Jr.,  La.,  19233,  pp.  36,  39). 

W.  S.  Blatchley  (Ind.,  1891,  p.  27)  calls  this  species  "the  most  abundant  tail- 
less batrachian  in  the  country.  Hundreds  are  to  be  seen  along  any  small  stream 
in  spring  and  autumn.  They  appear  less  common  in  summer." 

"This  species  was  the  most  abundant  amphibian  within  the  limits  of  this 
area.  It  was  found  most  abundantly  at  the  edges  of  bodies  of  water  in  all  parts 
of  the  area  in  spring  although  some  were  found  on  the  shores  of  the  lake 
through  summer  and  until  late  in  the  fall.  On  February  2,  1924,  several  were 
found  in  the  creek  above  the  bridge.  They  were  in  the  water  above  some  olcl 
ice  and  below  a  top  layer  of  new  ice.  All  the  frogs  were  stifl  and  floating  and 
apparently  they  were  dead.  In  the  warm  water  of  the  springs  and  just  below 
the  springs  a  few  frogs  of  this  species  were  found  throughout  the  winter"  (J.  M. 
Linsdale,  Kans.,  1927,  p.  76) . 

Size:  We  have  made  no  study  of  this  segregated  group.  A  tentative  summa- 
tion is:  adults,  %-i%  inches;  males,  17  or  18-30  mm.;  females,  20-35  mm.  Did 
Fowler's  (1907)  2%  inches  mean  i/4  inches? 

We  measured  20  males  and  20  females  from  Texas  and  Mississippi  Valley 
with  these  results:  males^  17.5-29.0  mm.,  average  21  mm.,  mean  23  mm.;  fe- 
males, 19-28  mm.,  average  24  mm.,  mean  23  mm.  In  one  lot  a  ig-mm.  male 


TREE  FROGS:  HYLIDAE  223 

was  debatable;  in  another  lot  three  individual  specimens,  17, 18, 18  mm.,  were 
hard  to  sex;  in  another  lot  we  had  some  males  17.5  and  18.5  mm. 

General  appearance:  In  1923  Viosca  (La.,  19233,  p.  43)  said:  "The  puzzling 
status  of  Acns,  as  far  as  Louisiana  is  concerned,  has  been  positively  cleared  by 
these  studies.  There  are  two  distinct  species  in  Louisiana,  the  upland  species 
being,  tentatively,  Acris  gryllus,  and  that  of  the  lowlands,  Acris  crepitans. 
Wherever  their  ranges  overlap,  they  are  found  side  by  side  without  inter- 
breeding, each  with  its  characteristic  chorus  and  habits."  Dunn  (Gen.,  1939, 
PP-  I53"I54)  confirmed  Viosca's  opinion  that  two  distinct  species  are  involved 
and  the  distinguishing  characters  are  those  he  mentioned:  "'The  best  char- 
acter for  distinguishing  the  two  species  is  the  amount  of  webbing  of  the  toes, 
crepitans  having  much  more  web.' 

"gryllus  crepitans 

Smaller  Larger 

Less  web  (5  phalanges  of  toe  More  web  (2  to  il/>  phalanges  of  toe 

4  free,  toe  i  partly  free)  4  free,  toe  i  completely  webbed) 

More  rugose  Smoother 

Anal  warts  less  prominent  Anal  warts  more  prominent 

Legs  longer,  heel  beyond  snout  Legs  shorter,  heel  not  to  snout 

Thigh  more  definitely  striped  Thigh  less  definitely  striped." 

"If  this  form  actually  exists,  as  Viosca  (1923,  1931)  maintains,  it  certainly 
does  not  exist  in  Kansas.  The  specimens  examined  are  too  uniform  in  charac- 
ter to  permit  of  more  than  one  species,  and  the  extent  of  variation  is  well 
within  that  of  gryllus'  (H.  M.  Smith,  Kans.,  1934,  p.  461). 

"Dunn  (Gen.,  1939,  p.  154)  lists  six  criteria  for  use  in  distinguishing  crept- 
tans  from  gryllus.  Four  of  these,  (i)  larger  size,  (2)  more  extensively  webbed 
feet,  (3)  shorter  legs,  and  (4)  less  definitely  striped  thighs,  are  evident  in  our 
specimens  when  they  are  compared  with  topotypes  of  gryllus  from  Riceboro, 
Georgia.  The  two  other  diagnostic  characters  used  by  Dunn  do  not  hold  in 
our  specimens;  they  seem,  therefore,  to  merit  further  discussion.  First  he  char- 
acterizes gryllus  as  'more  rugose'  than  crepitans;  the  contrary  is  true  in  the 
Rockingham  County  specimens  which  are  definitely  more  rugose  than  are 
gryllus  topotypes.  Wide  variation  in  the  amount  of  rugosity  occurs  in  both 
species  and  we  feel  that  this  character  is  the  least  useful  of  those  used  by 
Dunn.  Secondly,  Dunn  states  that  the  anal  warts  of  gryllus  are  less  prominent 
than  those  of  crepitans.  This  statement  is  ambiguous  because  it  fails  to  indi- 
cate whether  the  warts  are  prominent  by  reason  of  their  color,  size,  or  number. 
The  startlingly  white  color  of  the  subanal  warts  of  some  specimens  of  Acris 
is  a  result  of  preservation  in  formalin,  as  the  senior  author  has  determined  ex- 
perimentally; specimens  preserved  in  alcohol  show  less  color  change  and  the 
subanal  warts  do  not  fade  to  an  ivory  white.  It  is  quite  evident  however  that 
rugosity  of  the  central  thigh  area  is  more  characteristic  of  crepitans  than  of 


224  HANDBOOK  OF  FROGS  AND  TOADS 

gryllus;  the  Virginia  specimens  and  many  examples  of  crepitans  from  else- 
where exhibit  a  greater  number  of  subanal  warts  than  do  topotypes  of  gryllus, 
but  brilliant  white  warts  are  present  in  some  specimens  of  each  species.  In 
addition  to  the  characters  mentioned  by  Dunn,  crepitans  has  a  shorter  head 
and  a  more  obtuse  snout  than  gryllus"  (M.  G.  Netting  and  L.  W.  Wilson, 
W,Va.,  1940,  p.  6). 

Color:  "Color  above,  some  shade  of  gray,  brown,  or  olive-green,  often  with  a 
median  longitudinal  diffuse  band  of  red  or  green,  and  with  several  black  spots, 
of  which  a  triangular  one  between  the  eyes  is  constant  and  characteristic.  Be- 
neath pale.  Upper  jaw  black  or  dark  brown,  with  four  vertical  pale  lines  on 
each  side.  A  narrow  pale  line  extends  from  the  lower  posterior  part  of  the  eye 
to  the  base  of  the  fore  leg.  Above  this  line  lies  an  elongate  black  spot  which  ex- 
tends from  the  eye  towards,  but  does  not  quite  reach,  the  fore  leg.  Behind  the 
insertion  of  the  fore  leg,  on  the  side,  is  a  large  oblique  black  spot  margined  with 
white.  Another  similar  but  smaller  spot  lies  in  advance  of,  and  above,  the  in- 
sertion of  the  hind  leg.  The  triangular  spot  between  the  eyes  is  narrowly  mar- 
gined with  white,  its  apex  pointing  backward.  The  middle  of  the  back  is  often 
occupied  by  a  longitudinal  red  or  green  band,  and  immediately  on  each  side  of 
the  latter  are  several  obscure  black  spots.  Color  beneath  pale,  sometimes  tinged 
with  yellow  on  the  throat.  Throat  more  or  less  speckled  with  dusky  or  brown. 
Lower  jaw  pale,  or  with  a  few  dark  specks  at  the  symphysis,  becoming  darker 
towards  the  angle  of  the  mouth,  from  which  point  a  dark  dash  passes  to  and 
upon  the  base  of  the  fore  leg.  Legs  and  digits  dark  above,  with  round  dark 
spots;  pale  and  unmarked  below.  A  black  spot  may  often  be  visible  over  the 
vent,  and  generally  a  dark  bar  passes  from  this  region  along  the  posterior  sur- 
face of  the  thigh"  (H.  Carman,  111.,  1892,  p.  341). 

"Acris  gryllus.  Cricket  frogs  are  common  along  the  shores  of  all  bodies  of 
water.  Some  of  them  were  very  large.  All  were  dark  in  color  and  most  of  them 
showed  either  a  red  or  green  streak  down  the  center  of  the  back"  (K.  P. 
Schmidt,  III.,  1923,  p.  49) . 

Structure:  (See  A.  gryllus  gryllus  for  differences  in  the  two  forms.) 

"Width  of  head  across  base  of  lower  jaw  about  equal  to  distance  from  that 
point  forward  to  tip  of  snout;  legs  shorter  and  hind  feet  more  fully  webbeH, 
the  web  on  the  first  (shortest)  toe  reaching  its  tip;  heel  when  leg  is  extended 
not  reaching  tip  of  snout— Common  Cricket"  (C.  S.  Brimley,  N.C.,  1940, 
p.  15). 

"Webbing  on  toes  more  extensive;  last  two  phalanges  of  fourth  toe  free  from 
webbing;  usually  with  third  and  fifth  toes  reaching  beyond  middle  of  the  third 
phalanx  from  tip  of  fourth  toe.  A.  g.  crepitans"  (E.  B.  Chamberlain,  S.C.,  1939, 
p. 12). 

"None  of  this  questionable  species  was  found  in  Beaufort  County  although 
Brimley  (1926)  gives  the  range  in  North  Carolina  as  'central  and  part  of  east- 
ern districts.'  The  writer  has  seen  individuals  commonly  in  Durham  County 


TREE  FROGS:  HYLIDAE  225 

which  are  referable  to  this  species.  Viosca  (1923)  has  taken  considerable  inter- 
est in  the  genus,  as  a  result  of  field  observations  maintaining  that  in  Louisiana 
there  are  two  distinct  species  of  Acris.  The  writer  is  inclined  to  the  view  that 
more  field  and  experimental  study  is  needed  to  solve  the  problem.  It  was  ob- 
served that  in  Chase's  Lake,  Brooksville,  Mississippi,  only  typical  gryllus  forms 
occur  while  at  Cryme's  Pond  both  forms  occur  with  calls  which  are  recogniza- 
bly different.  The  former  locality  is  in  a  Lafayette  Red  Clay  area  while  the 
latter,  less  than  four  miles  distant,  is  in  a  Selma  Chalk  area"  (B.  B.  Brandt, 
N.C.,  19363,  p.  217). 

Voice:  Feb.  24,  1925,  Helotes,  Tex.  Tonight  air  64°  at  8:50  P.M.  Heard  at 
"Ornate  Fork"  quite  a  chorus  of  Acris,  gicl(,  gic^  gicf( — or  l(icl(,  %icl{,  ^icl{. 

March  8-12,  Helotes,  Tex.  Evenings — in  great  chorus  for  several  days.  Creek 
is  filled  with  them. 

"Besides  their  rattling  call  a  squeaky  sound  was  heard  occasionally,  though 
only  during  the  breeding  season.  .  .  . 

"It  may  be  stated  that  in  my  experience  their  call  appears  to  be  variable.  The 
usual  note  is  not  heard  at  a  great  distance,  and  is  described  by  one  writer  as 
exactly  imitated  by  striking  two  marbles  together,  first  slowly,  then  faster  and 
faster,  for  a  succession  of  about  20  to  30  beats.  Perhaps  the  rattling  of  castanets 
would  be  a  better  suggestion"  (H.  W.  Fowler,  N.J.,  1907,  pp.  102-103). 

"The  name  Cricket  Frog  was  given  to  it,  on  account  of  its  song,  which  bears 
a  strong  resemblance  to  the  chirping  of  the  black  cricket.  These  tiny  frogs  sing 
in  chorus  in  spring.  The  sound  can  be  imitated  by  striking  together  two  peb- 
bles or  two  marbles,  beginning  slowly  and  continuing  more  rapidly  for  thirty 
or  forty  strokes.  The  male  frog  is  the  singer  and  in  doing  so  inflates  his  yellow 
throat  enormously.  .  .  .  The  first  warm  days  in  early  spring  bring  them  out. 
Feb.  14,  20;  Mar.  5;  May  i;  Sept.  7;  Oct.  16"  (J.  Hurter,  Mo.,  1911,  p. 
102). 

"Feb.  8,  1918.  Buffalo  Bayou.  They  are  fairly  abundant  all  along  the  banks 
and  are  calling  vigorously.  The  call  is  a  soft  trill  resembling  the  tree  cricket  or 
the  mole  cricket"  (P.  H.  Pope,  Tex.,  1919,  p.  97). 

"The  cricket-like  chirping  of  these  abundant  little  frogs  is  a  familiar  sound 
about  the  marshy  borders  of  streams  and  ponds  from  April  to  July.  The  singer 
is  often  hard  to  locate  due  to  the  carrying  capacity  of  the  sound  and  a  more  or 
less  ventriloqual  effect"  (G.  E.  Hudson,  Neb.,  1942,  p.  25). 

"The  voice  of  a  cricket  frog  is  a  combination  of  a  rattle  and  a  musical  clink, 
but  it  is  only  about  half  as  loud  as  that  of  a  spring  peeper.  A  chorus  heard  at  a 
distance  sounds  like  the  jingling  of  small  sleigh  bells,  for  the  musical  element 
of  its  call  travels  farther  than  the  rattle.  A  chorus  heard  close  by  sounds  like  the 
rattle  of  small  pebbles  poured  upon  a  cement  pavement. 

"An  individual  frog  sings  for  from  thirty  to  forty-five  seconds  at  a  time.  Its 
call  has  three  phases.  The  first  phase  lasts  for  about  five  seconds  and  sounds  like 
the  clicks  of  a  boy's  marble  dropped  upon  a  cement  pavement,  once  or  twice  a 


226  HANDBOOK  OF  FROGS  AND  TOADS 

second  from  a  height  of  about  six  inches.  The  second  phase  sounds  like  the  gal- 
loping of  a  small  pony  on  a  brick  pavement,  or  like  the  clicks  of  a  boy's  marble 
dropped  upon  a  pavement  from  a  height  of  only  an  inch  or  two,  and  allowed 
to  bounce  each  time.  The  third  phase  sounds  like  the  regular  cree-cree-creeing 
of  a  tree  cricket,  or  like  the  rattle  of  a  boy's  marble  that  bounces  rapidly  when 
it  is  dropped  at  frequent  intervals  from  a  height  of  only  half  an  inch.  The  time 
and  rhythm  of  the  sounds  are  about  the  same  as  that  of  the  following  syllables 
pronounced  with  the  speed  of  ordinary  reading:  click,  click,  click,  click 
.  .  .  click-e-ty,  click-e-ty,  click-e-ty,  click-e-ty,  click-e-ty  .  .  .  cree,  cree,  cree, 
cree.  .  .  . 

"The  cricket  frog  inflates  a  vocal  sac  under  its  chin  during  its  call.  It  often 
sits  quietly  with  its  sac  distended  for  many  minutes  between  its  calls.  The  vio- 
lent efforts  of  its  body  in  producing  its  sound  make  the  frog  resemble  a  small 
boy  on  his  hands  and  knees  blowing  a  fire  with  all  his  might.  The  vocal  sac  is 
bright  yellow,  and  when  it  is  seen  distended  in  the  day  time,  it  is  so  conspicuous 
that  it  reveals  many  a  singer  that  otherwise  would  be  almost  invisible  on  a  lily 
pad"  (F.  Overton,  N.Y.,  1914,  p.  31). 

Breeding:  Early  April-July.  (See  A.  g.  gryllus.)  H.  M.  Smith  (Kans.,  1934, 
p.  389)  gives  the  egg  as  having  a  single  envelope  2.3  mm.  or  more.  Dr.  Kather- 
ine  Van  Winkle  (Mrs.  E.  L.)  Palmer  who  worked  over  our  Acns  collections 
and  our  1917  transcontinental  material  found  that  in  an  Ames,  Iowa,  congress 
of  Acris  June  12,  1926,  some  eggs  tended  to  be  in  bunches  or  masses.  We  meas- 
ured her  Ames  material  and  found  males  21-29  mrn-  m  length  and  females  27- 
29  mm.  Most  of  the  records  indicate  transformation  from  13-15  mm.,  but  we 
were  quizzical  and,  on  testing  out  our  Texas  material,  found  they  ran  mainly 
from  1T-I2  mm.,  sometimes  10  or  10.5  and  12.5  or  13  mm.  An  inclusive  range 
of  10-15  mm.  is,  therefore,  better  for  transformation  sizes.  In  the  same  way, 
few  tadpoles  reach  45  or  46  mm.;  most  run  from  30  to  36  mm.,  with  the  body  of 
the  tadpole  usually  11.5-14.0  mm. 

Miss  Dickerson  believed  their  chorus  was  loudest  in  late  April  and  early  May 
when  they  attached  their  eggs  to  grass  blades  or  leaves  in  the  water.  She  stated: 
"The  development  of  this  frog  is  less  rapid  than  that  of  the  Common  Tree 
Frog,  the  Eastern  Wood  Frog,  or  the  American  Toad.  The  tadpoles  may  be 
found  in  the  water  as  late  as  August.  The  final  transformation  takes  place  in 
September.  The  young  tree  frogs,  as  well  as  the  older  ones,  seek  shelter  from 
the  cold  under  stones  and  leaves  at  the  margins  of  their  brook  or  marsh." 

"[Eggs  are]  not  deposited  in  large  groups,  and  not  concealed  under  objects; 
outer  envelope  usually  not  over  7.5  cm.  Envelope  single.  Envelope  2.3  mm.  or 
more — Acris  gryllus  (Le  Conte)"  (H.  M.  Smith,  Kans.,  1934,  p.  389). 

"Although  individuals  are  sometimes  collected  throughout  the  winter,  and 
are  rather  abundant  in  March,  their  breeding  activities  do  not  begin  until  early 
April.  .  .  .  Near  Lawrence  clasping  pairs  have  been  collected  as  late  as  May  9 
(1933) ;  they  probably  breed  much  later,  as  their  songs  are  frequently  heard  as 
late  as  July,  and  Gloyd  (personal  notes)  has  heard  them  singing  about  Man- 


TREE  FROGS:  HYLTDAE  227 

hattan  as  late  as  July  15.  Eggs  have  been  laid  in  the  laboratory  on  May  10,  from 
females  collected  the  preceding  night  (1933).  The  breeding  sites  chosen  are 
varied.  In  regions  where  the  species  is  found  in  abundance,  permanent  lakes, 
streams  and  springs  always  have  their  quota.  Frequently  they  breed  in  and  sing 
from  temporary  pools  in  pastures  or  at  road-sides"  (same,  p.  459). 

"On  July  17,  1932,  metamorphosis  was  occurring  in  these  frogs  at  Isle  Du 
Bois  Creek,  where,  it  was  estimated,  many  thousands  of  these  small  frogs  oc- 
curred, and  where  specimens  in  almost  all  stages  of  development  were  present. 
Some  juveniles  without  fully  absorbed  tails  had  bright  green  dorsal  patterns, 
while  tailed  specimens  approaching  transformation  possessed  the  typical  tri- 
angular dorsal  marking,  but  no  such  larvae  were  observed  to  have  the  black 
tail  tip  characteristic  of  the  tadpoles  occurring  in  a  woodland  pond  at  Danby, 
about  two  miles  distant.  A  transforming  specimen  upon  emergence  from  the 
water  on  August  9  possessed  a  body  length  of  14.4  mm.,  tail  14.9.  Two  hours 
and  forty  minutes  later  the  tail  measurement  was  10.75,  an  absorption  of  4.15 
during  this  period.  Another  larva,  measured  at  9:15  P.M.,  had  a  body  length  of 
15.1,  tail  31.7,  of  which  9  mm.  possessed  dark  pigmentation;  the  mouth  was 
transformed,  the  dorsal  pattern  distinct.  By  7:30  P.M.  August  10  the  body 
length  was  14.4,  tail,  24,  dark  tip,  3.5  and  by  2  P.M.  August  u  the  body 
length  was  14,  tail,  1.5.  In  this  period  of  40%  hours  the  reduction  in  body 
length  was  i.i  mm.,  tail  30.2.  By  September  18  the  transformation  period  was 
neanng  its  end  and  few  larvae  could  be  taken  by  seining  at  this  time.  On  this 
date,  however,  a  ribbon  snake,  Thamnophis  sattnttts  proxtmns  was  lying  on 
a  mat  of  aquatic  vegetation  at  a  pond  near  Danby,  and  when  captured  it  dis- 
gorged three  cricket  frogs,  one  a  transforming  specimen"  (D.  A.  Boycr  and 
A.  A.  Hemze,  Mo.,  1934,  P-  x<^9)' 

"The  eggs  were  laid  singly  or  in  small  masses.  One  female  had  248  .  .  ." 
(G.  E.  Hudson,  Neb.,  1942,  p.  25). 

Journal  notes:  We  have  seen  it  from  New  Jersey  to  Georgia,  Louisiana, 
Texas,  and  Iowa,  but  like  most  workers  we  have  neglected  this  most  ubiqui- 
tous tree  frog.  Everyone  remarks  their  choruses.  We  will  forbear  except  to 
give  the  scanty  notes  of  two  seasons  in  Texas. 

Feb.  8, 1925,  Rio  Cibola  River,  Tex.  A  few  Acns  called. 

Feb.  8-Feb.  21,  Helotes.  Once  in  a  while  hear  a  few  Acns  calling.  Not  a  loud 
full-throated  Acrls  call  yet. 

Feb.  21.  Heard  them  calling,  air  and  water  7o°F.,  9-10  P.M. 

Feb.  24  to  March  8-12.  [See  "Voice."  | 

March  13.  In  great  chorus  last  night  and  tonight.  In  our  fork  it  is  very 
common. 

March  16.  Last  night  no  Acris  calling. 

April  i.  Heard  cricket  frogs  at  Sabinal,  at  Nueces  River,  and  at  Uvalde. 

April  2,  Cotulla.  Heard  in  Nueces  River  or  extension  many,  many  Acris. 

May  5,  Helotes.  On  talus  slope  we  turned  over  a  flat  stone  and  saw  a  greenish 
frog.  Another  jumped  and  my  eye  followed  it.  It  impressed  me  as  Acris.  I  let 


228  HANDBOOK  OF  FROGS  AND  TOADS 

the  stone  down.  Later  looked  under  the  stone,  and  behold  a  Marnock's  frog — 
a  red-letter  day.  Later  we  went  to  the  spot  where  the  other  frog  jumped  and 
it  proved  to  be  an  Acris.  It  must  have  finished  breeding  to  be  here. 

May  9,  San  Antonio,  8:30  P.M.  Acris  calling  strongly  near  St.  Mary's  College 
and  in  Bandera. 

May  14,  Boerne.  Heard  a  few  Acris. 

May  24,  Helotes.  Went  to  Lee's  Branch  to  get  Acris  crepitans  series.  Many 
Acris  transformed. 

June  6,  Devine.  Found  Acris  tadpoles  with  black-tipped  tails. 

Jan.  17,  1942,  With  Quillms  to  Classen  Ranch.  In  clear  Fork  of  Cibola  .  .  . 
Roy  dug  under  roots  of  a  tree  and  brought  out  a  Bufo  valliceps.  In  the  creek 
saw  plenty  of  Acris  crepitans,  tadpoles  minute  to  almost  mature  of  R.  pipiens. 
Saw  an  immense  R.  catesbeiana  male  in  mass  of  vegetation. 

July  22,  Toyahvale,  Tex.  Tonight  at  9:30  drove  to  where  Cherry  Canyon 
goes  across  the  highway  and  several  miles  beyond.  At  first  we  heard  a  note 
1  thought  was  a  rattle  in  rear  of  our  auto,  but  Anna  persisted,  and  then  I  got  it: 
full  Acris  crepitans.  Got  plenty  of  them  for  first  2  or  3  miles  west  of  Toyahvale. 

July  23.  Trip  to  Phantom  Lake  with  Alex  Izzard  of  Balmorhca  State  Park. 
Saw  several  Acns  here  and  caught  them.  Water  about  70 °F.  A  fine  spring. 

August  2.  Beyond  Carrizo  Springs,  Tex.  At  Sycamore  Cr.  boundary  line  of 
two  counties  took  one  transformed  Acris. 

In  Texas  in  1925  we  began  trips  about  February  8,  when  Acris  appeared  if 
nothing  else.  For  many  entries  we  merely  say,  "No  end  of  Acris"  "Acris  in  all 
creeks  of  whatever  size."  Always  merely  "Acris."  Not  until  May  24-26,  when 
they  were  transforming  and  transformed,  did  we  courageously  enter  A.  crepi- 
tans. At  present  we  are  almost  willing  to  grant  crepitans  and  gryllus,  but  are 
there  one  to  three  or  more  forms  of  Acris  yet  undescribed  ?  This  omnipresent 
species  may  prove  more  involved  than  Rana  pipiens. 

Authorities9  corner: 

H.  Garman,  111.,  1892,  p.  342.  C.  S.  Brimley,  N.C.,  1940,  p.  15. 

J.  Hurter,  Mo.,  1911,  p.  101.  G.  A.  Moore  and  C.  C.  Rigney,  Okla., 

T.  L.  Hankinson,  111.,  1917,  p.  324.  1942,  p.  78. 

B.  W.  Evermann  and  H.  W.  Clark,     G.  L.  Orton,  La.,  1947,  pp.  375-377. 

Ind.,  1920,  p.  634. 

In  a  recent  paper  (May  10,  1947)  F.  Harper  (Gen.,  1947)  restricts  Acris  g. 
crepitans  to  the  eastern  portion  of  its  credited  range,  i.e.,  from  Connecticut  to 
New  York  to  eastern  Texas,  and  establishes  a  new  form,  A.  g.  blanchardi  for 
the  cricket  frogs  from  southwestern  Michigan  to  southwestern  Missouri  to 
southwestern  Texas  and  westward  to  Utah  and  Arizona.  The  new  form  "is 
distinguished  ...  by  slightly  greater  linear  measurements,  by  decidedly 
greater  bulk,  by  somewhat  more  extensive  dusky  area  on  the  posterior  face  of 
the  femora  in  the  vicinity  of  the  vent."  Our  preceding  discussion  of  Acris  g. 
crepitans  is  therefore  composite  if  this  interpretation  proves  the  correct  one. 


Genus  PSEUD ACRIS  Fitzinger l 
Maps  1 6-1 8 

Mountain  Chorus  Frog,  Chorus  Frog,  Ohio  Chorus  Frog 
Pseudacrts  brachyphona  (Cope).  Plate  XLVII;  Map  16. 

Range:  Southwestern  Pennsylvania,  western  Maryland,  southeastern  Ohio, 
eastern  Kentucky,  West  Virginia,  northern  Alabama  (Viosca),  northern  Mis- 
sissippi. Must  be  in  Tennessee. 

Habitat:  Springy  hillsides,  grassy  pools,  ditches,  sources,  and  along  upper 
courses  of  upland  rivulets — more  hilly  than  lowland  habitats. 

Size:  Adults,  1-1%  inches.  Males,  24-52  mm.  Females,  27-34  mm- 

General  appearance:  These  arc  small  frogs,  gray  or  brown  in  color,  medium 
in  size  for  Pseudacns,  with  the  most  distinct  digital  disks  of  this  genus.  They 
arc  more  stocky  in  body  and  broader  in  head  than  P.  n.  triseriata  and  P.  n. 
feriarum.  The  usual  middorsal  stripe  or  row  of  spots  is  lacking.  They  often 
have  a  light  middorsal  area  somewhat  after  the  pattern  of  the  cricket  frog 
(Acris  gryllus).  The  intcrorbital  triangle  is  not  white-edged  behind.  The  dor- 
solateral  bands  curve  from  the  eye  to  midback  to  groin,  making  two  crescents. 
They  often  meet  in  midback  to  form  a  cross  or  transverse  bar.  Sometimes  the 
pattern  consists  of  a  cross  or  a  bar  alone. 

Color:  (See  "General  appearance,"  "Structure,"  and  "Authorities'  corner.") 
In  general  color  these  frogs  range  from  the  sorghum  brown,  deep  brownish 
drab,  or  mars  brown  of  Hyla  femoralis  to  some  of  the  grays  found  in  Acris  or 
Hyla  femoralis  or  to  the  blackish  olive  of  Pseudacns  n.  triseriata. 

Structure:  The  original  description— "A  specimen  of  nearly  the  si/.e  and 
form  of  Hyla  femoralis  was  taken  in  western  Pennsylvania,  near  the  Kiskimi- 
nitas  River.  In  proportions  it  does  not  differ  from  the  Feriarum,  but  the  toes 
arc  fringed,  the  dilations  larger  and  the  coloration  different.  Above  blackish- 
ash,  abruptly  defined  on  the  sides.  Lateral  band  not  extending  beyond  tympa- 
num. No  median  dorsal  band,  but  two  black  dorso-laterals  of  double  ordinary 
width  converge  from  each  tympanum  and  extend  to  end  of  urostyle  inclosing 
with  the  interorbital  triangle  a  narrow,  anteriorly  bifurcate  dorsal  band  of 
ground  color"  (E.  D.  Cope,  Gen.,  1889,  p.  341). 

Compared  to  P.  ornata,  P.  n.  feriarum,  P.  n.  triseriata,  it  has  a  wider  head 

1  The  two  workers  who  in  the  last  ten  to  fifteen  years  have  added  most  to  clarification 
of  this  genus  are  C.  Walker  and  F.  Harper. 


230 


HANDBOOK  OF  FROGS  AND  TOADS 


and  a  longer  hind  limb,  tibia,  and  foot  with  tarsus.  In  most  ways  it  falls  into 
the  Pseudacris  ornata  group;  in  some  with  the  Pseudacris  nignta  group,  e.g., 
as  shown  in  the  statistics  (number  of  times  in  length)  below: 

P.  ornata    P.  brachyphona    P.  n.  feriarum 
Width 
of  head  2.86-3.  2.5-3.1 

0.60-0.67 

i.  82-  r.  9  3 


P.  n.  triseriata 


Hind  limb 
Tibia 
Foot  with 
tarsus 


2.86-3.4 

0.68-0.75 

2.06-2.28 


3-0-3-33 

0.65-0.74 

2.0-2.26 


3-25-3-5 

0.71-0.75 

2.24-2.54 


1.26-1.42 


1.23-1.60 


I-37-I-57 


These  measurements  distinguish  it  from  P.  n.  triseriata,  but  not  so  clearly  from 
P.  n.  feriarum.  The  proportions  for  P.  brachyphona  are  in  three  parts  very  close 


Pseudacris  brachyphona 


16 


to  those  of  P.  n.  nigrita.  Although  we  have  put  this  species  in  the  P.  ornata 
group  in  the  key,  nevertheless  there  are  some  respects  in  which  it  is  not  clearly 
divorced  from  the  P.  nigrita  six  subspecies. 

Voice:  "The  note  of  this  species  is  quite  different  from  that  of  the  C.  triseria- 
ttts,  not  being  continuous,  but  in  sets  of  crepitations  repeated  in  time  and  at  in- 
tervals" (E.  D.  Cope,  Gen.,  1889,  p.  341).  When  P.  n.  triseriata  and  P.  bra- 
chyphona are  calling  they  are  different;  the  latter  has  a  faster,  higher  note,  yet 
its  call  belongs  distinctly  in  quality  and  form  with  the  P.  nigrita  group. 


TREE  FROGS:  HYLIDAE  231 

"The  voices  of  the  two  (tnsenatits  and  brachyphona)  are  much  alike  but  the 
call  of  brachyphona  is  given  more  quickly,  with  a  higher  pitch  and  a  different 
quality  so  that  the  effect  of  a  chorus  is  quite  distinctive"  (C.  F.  Walker,  Ohio, 


"Their  notes  are  repeated  at  the  rate  of  about  50  to  70  times  a  minute  and 
may  be  continued  for  several  minutes  although  they  usually  stop  in  fifteen  to 
twenty  seconds.  The  call  is  strong  and  rapid  and  on  a  clear  night  a  chorus  may 
carry  a  quarter  of  a  mile"  (N.  B.  Green,  W.  Va.,  1938,  pp.  79-80). 

Breeding:  March-July.  Amplexation  axillary.  Eggs  in  masses,  10-50,  at- 
tached to  vegetation  or  trash.  Egg  complement  300-1500;  vitellus  1.6  mm.;  en- 
velope 6.0-8.5  mm.  Tadpole  period  50-60  days.  Tadpole  25  mm.  Labial  tooth 
rows  %.  Transformation  size  8.0  mm.  (after  Green,  Walker,  Welter,  Carr, 
Barbour,  and  Sollberger,  the  last  four  of  whom  have  sent  us  many  a  puzzling 
specimen  of  this  species,  ot  Hyla  cructfer,  and  of  Acns). 

"Its  breeding  season  begins  somewhat  later  than  that  of  trisenata  and  is  more 
prolonged.  Clasping  pairs  and  fresh  eggs  have  been  found  as  early  as  March 
2oth  and  as  late  as  May  i2th.  In  the  same  region  our  dates  for  tnsenata  eggs 
range  from  March  i4th  to  April  i6th.  The  eggs  of  brachyphona  are  so  much 
like  those  of  tnsenata  and  arc  laid  in  similar  masses,  attached  to  twigs,  leaves, 
or  grasses,  often  well  below  the  surface  of  the  water"  (C.  F.  Walker,  Ohio, 

«932,  P-  383)- 

"Egg  masses  were  found  in  the  tegion  under  observation  (Randolph  and 
Tucker  counties,  West  Virginia,  at  an  elevation  of  2000  feet)  on  April  5. 
Walker  (1932)  states  that  iresh  eggs  have  been  found  as  early  as  March  20, 
presumably  in  southern  Ohio.  Pairs  in  amplcxus  and  eggs  were  taken  at  Por- 
terwood,  Tucker  County,  on  June  10,  1936,  at  an  elevation  of  2200  feet  and  on 
Point  Mt.,  Randolph  County,  on  June  7,  1937,  at  an  elevation  of  3500  feet  fol- 
lowing a  rainy  spell  of  several  days.  Freshly  laid  eggs  were  collected  by  C.  J. 
Coin  at  Camp  Woodbine,  Nicholas  County,  on  July  2,  1936,  at  an  elevation  of 
2000  feet.  His  notes  for  this  date  read,  'Small  pool  visited  at  dusk  following  a 
rain.  A  long  dry  spell  had  preceded  this  rain.  One  clutch  of  freshly  deposited 
eggs  was  observed.  .  .  .'  On  April  13,  1937,  five  females  were  collected  on  their 
way  to  the  pools.  In  the  laboratory  each  female  was  put  in  a  separate  jar  with 
a  male  in  an  effort  to  determine  the  number  of  eggs  an  individual  laid.  They 
laid  respectively  1479,  383,  318,  and  406  eggs"  (N.  B.  Green,  W.  Va.,  19^8, 
p.  80)  . 

"Egg  masses  were  collected  from  a  conical  hole  about  two  feet  deep  in  the 
vicinity  of  Reightown  on  April  22,  1939.  On  the  evening  of  June  30,  1939  a 
single  specimen  was  collected  in  a  ditch  in  a  field  adjacent  to  the  highway  be- 
tween Homer  and  Ruggles  Gap"  (H.  D.  Yoder,  Pa.,  1940,  p.  92). 

"A  very  common  spring  frog.  Breeding  begins  as  early  as  February  12  when 
the  marshes  ring  with  the  songs  of  this  species  and  of  Hyla  cructfer.  It  is  as 
abundant  as  H.  critcifer  and  is  usually  found  with  it"  (W.  A.  Welter  and  K. 
Carr,Ky.,  1939,?.  129). 


231  HANDBOOK  OF  FROGS  AND  TOADS 

Journal  notes:  July  18,  1931,  Beckley,  W.  Va.,  Fair  Grounds  woods.  The 
botanists  found  a  queer  Psettdacns  jenantm  near  a  sawdust  pile. 

July  19,  Beckley,  W.  Va.,  Fair  Grounds.  Tonight  after  a  heavy  ram,  in  a 
puddle  beside  the  grandstand  and  in  pools  beside  the  track  heard  several  Hyla 
versicolor,  and  in  the  distance  several  Hyla  crucijer,  but  the  notes  which  drew 
us  to  the  spot  were  new — surely  a  Pseudacris  in  character  of  voice,  not  a  Hyla 
crucijer  nor  an  Acris.  It  was  different  from  Pseudacris  n.  tnscriata  and  Pseuda- 
cns  n.  jenantm.  When  we  first  caught  it,  its  long  legs  reminded  us  of  an  Acris, 
and  we  looked  for  the  rear  femoral  stripes.  They  were  not  present.  It  seems  like 
a  Pseudacris,  but  has  too  large  disks  for  a  normal  Pseudacris  species.  Must  look 
up  Cope's  P.  j.  brachyphona. 

July  21.  Prof.  P.  C.  Bibbee,  of  State  College  at  Athens,  Mercer  Co.,  brought 
us  a  jar  of  preserved  swamp  cricket  frogs.  They  have  such  queer  coloration 
(more  like  an  Acns  type)  and  such  spindling  legs.  Are  they  a  Hyla,  Pseudacris, 
or  Acris?  They  must  be  Pseudacris.  Bibbee  let  me  take  a  few  for  our  collection 
and  said  they  collect  them  in  the  spring.  Surely  someone  will  have  to  collect 
Pseudacris  across  southern  Pennsylvania  from  Carlisle  (type  locality  of  P.  jeria- 
rum)  to  southwestern  Pennsylvania  and  West  Virginia  to  see  where  this  form, 
if  P.  f.  brachyphona,  meets  P.  fenarum  jenantm  9  m  mid-Maryland  or  mid- 
Pennsylvania. 

July  24.  Visited  R.  K.  Brown's  laboratory  at  Blucficld,  W.  Va.  He  had  a  jar 
with  several  puzzling  Pseudacris. 

C.  F.  Walker  was  the  first  author  to  redescribe  Cope's  Chorophilus  jenarum 
brachyphonust  and  it  is  interesting  that  four  groups  at  about  the  same  time 
independently  concluded  to  revive  Cope's  description;  Walker  on  Ohio  ma- 
terial, Netting  on  west  Pennsylvania  and  northern  West  Virginia  material, 
Dunn  on  west  Maryland  and  Pennsylvania  specimens,  and  we  on  southern 
West  Virginia  material.  But  the  question  is  not  entirely  settled  yet.  Is  this  form 
in  eastern  Tennessee ?  Even  as  early  as  1931  or  1933  when  this  form  was  re- 
vived, we  had  seen  material  from  northern  Mississippi  and  northern  Alabama 
to  cause  us  to  credit  it  to  those  states.  Viosca  (Ala.,  1938,  p.  201)  furnished  a 
very  fortunate  record. 

A  former  student  supplies  the  following:  "The  surprise  of  the  trip  was  a 
voice  heard  for  the  first  time  near  Moundville.  The  origin  of  the  strange 
'Qurack,'  suggestive  of  the  voice  of  Hyla  squirella,  eluded  us  for  about  twc 
hours  before  the  first  specimen  was  located;  then  the  next  six,  all  males 
were  taken  in  about  15  minutes.  This  locality,  in  the  foothills  at  the  extreme 
southwestern  limits  of  the  Appalachian  Highlands,  extends  the  range  of  this 
species  \P.  brachyphona}  some  500  miles  southwestward"  (letter).  "To  deter 
mine  the  number  of  eggs  laid  by  each  female,  ten  average-sized  individuals 
were  dissected.  The  largest  number  of  eggs  found  in  any  one  of  these  was  1202 
and  the  smallest  number  was  983.  The  average  number  found  in  ten  female* 
was  1092  eggs.  The  eggs  were  deposited  in  masses  attached  to  a  stem  or  leaf. 


TREE  FROGS:  HYLIDAE 


Plate  XLV1I.   Pxudacris  brachyphona  Plate  XLVIII.  Pseudacris  brimlcyt  (X 

XO.  1-7-  Males.  i,2,4,5>6-  Males-  3,7-  Females. 


234  HANDBOOK  OF  FROGS  AND  TOADS 

The  number  of  eggs  found  in  each  of  eleven  masses  was  counted,  and  the  num- 
ber found  to  vary  from  28  to  40,  the  average  being  34  per  mass.  In  the  labora- 
tory at  temperatures  varying  from  1 8  to  22  degrees  [C|,  hatching  occurred 
from  72  to  96  hours  after  the  eggs  were  laid.  In  the  pools,  however,  where  the 
temperature  ranged  from  2  to  13  degrees,  hatching  did  not  occur  for  a  week  or 
ten  daysS.  Several  tadpoles  kept  in  a  large  aquarium  in  the  laboratory  at  room 
temperature  metamorphosed  from  forty  to  fifty  days  after  hatching.  Tadpoles 
in  outdoor  pools  metamorphosed  in  forty-five  to  fifty-five  days"  (R.  W.  Bar- 
bour  and  E.  P.  Walters,  Ky.,  1941,  p.  116). 

Authorities9  corner: 
C.  F.  Walker,  Ohio,  1932,  pp.  380-381. 
P.  Viosca,  Jr.,  Ala.,  1938,  p.  201. 
N.  B.  Green,  W.  Va.,  1938,  p.  79. 

Brimley's  Chorus  Frog 

Pseudacns  brimleyi  Brandt  and  Walker.  Plate  XLVIII;  Map  16. 

Range:  Dismal  Swamp,  Va.,  south  in  the  Coastal  Plain  to  Bryan  Co.,  Ga.  In 
North  Carolina,  Brandt  has  taken  bnmleyi  in  Beaufort,  Greene,  Pitt,  and 
Wilson  counties,  and  has  voice  records  from  Craven  and  Edgecomb  counties. 
Newbern,  N.C.  Type  locality— Washington,  N.C.  (after  Brandt  and  Walker, 
N.C.,  1933,  p.  5).  Charleston,  S.C.  (E.  B.  Chamberlain,  S.C.,  1939,  p.  17). 

Habitat:  Heavily  wooded  flood-plain  pools.  Swampy  areas.  Brandt  (N.C., 
J93^b,  p.  500)  called  it  a  "palustrine  species.  .  .  .  This  chorus  frog  was  selected 
as  the  representative  of  an  arboreal  form  which  has  reverted  to  terrestrial  life." 
"It  is  of  especial  interest  in  comparison  with  its  more  arboreal  relative  of  simi- 
lar size,  Hyla  crncifer.  About  four  months  are  spent  at  the  breeding  pools;  the 
remainder  of  the  year  this  species  usually  spends  along  swampy  shores." 

Size:  Adults,  i-iVi  inches.  Males,  24-28  mm.  Females,  27.0-32.5  mm. 

General  appearance:  "A  dark  middorsal  stripe  from  tip  of  snout  to  vent.  On 
each  side  of  this  a  similar  stripe  extending  from  eyelid  to  posterior  end  of 
trunk.  A  darker,  sharply  defined  line  from  snout  through  eye,  and  along  side 
to  groin.  Dorsal  surfaces  of  legs  with  several  large  dark  spots,  the  long  axes  of 
which  are  longitudinal,  and  many  smaller  dark  spots.  A  distinct  dark  line 
along  outer  edge  of  tibia  and  on  under  side  of  foot.  A  light  line  along  upper 
jaw  continuing  back  to  shoulder.  Edge  of  jaw  dark.  An  irregular,  more  or  less 
discontinuous,  dark  line  along  posterior  side  of  arm.  Under  surface  light  in 
color  with  dark  spots  on  chest.  Smaller  but  similar  spots  on  under  surface  of 
legs"  (B.  B.  Brandt  and  C.  F.  Walker,  N.C.,  1033,  P-  3)  • 

"A  medium-sized  Pscndaeris  with  long  legs  and  a  trheriate  dorsal  pattern, 
closely  related  to  the  mgritajerianim-tnseriata-brachyphona  series,  but  differ- 
ing in  having  a  more  delicate,  smoother,  skin;  a  sharply  defined  black  lateral 
stripe  combined  with  a  rather  weak  dorsal  pattern;  a  pronounced  tendency 


TREE  FROGS:  HYLIDAE  235 

towards  longitudinal  rather  than  transverse  leg  markings;  a  narrow  dark  line 
along  outer  edge  of  tibia;  well  defined  dark  spots  on  chest  in  most  individuals; 
and  in  life,  a  distinctly  yellowish  venter"  (same,  pp.  2-3). 

Color:  "Dorsal  color  brown  (yellowish  to  reddish  shade) ;  dark  mid-dorsal 
stripe  from  snout  to  vent,  and  on  each  side  of  this  a  similar  stripe  from  eyelid 
to  posterior  end  of  trunk.  A  darker,  sharply  defined  lateral  line  from  snout 
through  eye  to  groin.  (In  light  phases  the  dorsal  stripes  are  usually  obscure,  but 
the  lateral  stripes  remain  black  and  sharply  defined.)  Under  surface  yellowish 
with  small  dark  spots"  (E.  B.  Chamberlain,  S.C.,  1939,  p.  17). 

"Living  specimens  show  considerable  variation  in  the  ground  color  of  the 
upper  parts,  but  are  apparently  restricted  to  yellowish  and  reddish  shades  of 
brown,  ranging  from  a  pale  buffy  brown  to  a  very  dark  shade  near  Light  Seal 
Brown  (Ridgeway).  The  dorsal  stripes  and  the  peculiarly  shaped  leg  spots  are 
usually  obscure  in  the  lighter  phases,  the  lateral  stripes  remain  black  and 
sharply  defined.  The  color  of  the  venter  also  varies  but  seems  always  to  be 
yellower  than  that  of  mgrtta.  The  iridescent  surface  makes  an  accurate  descrip- 
tion of  this  color  difficult.  In  the  paler  individuals  it  approaches  Cream-buff, 
and  in  more  richly  colored  examples  it  is  near  Buff-yellow  or  Pale  Orange- 
yellow.  The  light  cheek  stripe  below  the  vitta  is  similarly  colored.  Frequently 
there  is  a  deeper  yellow  spot  in  the  groin.  The  undersurface  of  the  legs  is  gray- 
ish. The  scattered  dusky  pigment  of  the  gular  sac  is  underlain  with  greenish 
yellow.  There  is  an  obscure  dark  bar  through  the  eye.  The  iris  shows  gold  and 
reddish  flecks,  the  latter  restricted  to  the  lower  half"  (B.  B.  Brandt  and  C.  F. 
Walker,  N.C.  and  Ga.,  1933,  p.  4). 

"The  more  distinctive  marks  arc  the  black  lateral  stripe  which  extends 
through  the  eye  almost  to  the  groin,  the  longitudinal  not  transverse  markings 
on  the  hind  legs,  and  the  spotted  under  parts"  (C.  S.  Brimlcy,  N.C.,  1940,  p.  18) . 

Structure:  Smoother  than  P.  n.  ntgrita.  Venter  coarsely  granulated;  heel  to 
anterior  eye;  disks  slightly  dilated;  muzzle  rounded  and  projecting;  tympa- 
num distinct,  smaller  than  eye. 

Brimley  (p.  15)  gives  these  characters:  "Upper  parts  with  small  warts  at  least 
on  the  side;  snout  longer  than  width  across  lower  jaw  at  base;  snout  rounded 
when  viewed  from  above;  breast  spotted." 

"The  leg  markings  of  brimleyt  and  oculans  remain  quite  different.  An  ap- 
proach to  the  longitudinal  spots  of  bnmleyi  is  seen  occasionally  in  brachy- 
phona,  and  the  voice  also  resembles  that  of  brachyphona,  which  is  a  very  dif- 
ferent frog  in  head  shape,  pattern,  and  skin  texture"  (B.  B.  Brandt  and  C.  L. 
Walker,  N.C.  and  Ga.,  1933,  p.  6) . 

Voice:  "Walker,  who  heard  the  species  in  northern  Georgia,  considers  the 
call  very  similar  to  that  of  Pscudacris  brachyphona.  To  the  writer  the  call 
sounds  like  a  rasping  trill,  'Kr-r-r-ak/  somewhat  less  than  a  second  in  duration. 
The  note  is  suggestive  of  that  of  Hyla  squirella  but  is  more  strongly  accented 
at  the  end,  and  the  intervals  between  the  individual  calls  are  shorter.  Calling 


236  HANDBOOK  OF  FROGS  AND  TOADS 

has  been  heard  in  rare  instances  as  early  as  November"  (B.  B.  Brandt,  N.C., 
19363,  p.  218). 

"The  type  locality  is  Washington,  N.C.,  where  it  is  the  earliest  breeding 
species  of  the  genus,  mated  pairs  having  been  taken  by  Brandt  and  his  friends 
of  the  Washington  Field  Museum  from  February  19  to  April  17, 1933"  (C.  S. 
Brimley,  N.C.,  1940,  p.  18). 

We  still  need  detailed  descriptions  of  the  egg  masses,  individual  eggs,  tad- 
poles, and  transformation  stages. 

Journal  notes:  Feb.  22, 1934.  We  have  visited  the  Bughouse  at  Washington, 
N.C.,  and  George  Ross  has  shown  us  some  of  the  live  adults  and  tadpoles  of 
P.  brimleyt.  There  are  yet  no  detailed  descriptions  of  the  eggs  and  tadpoles, 
but  they  do  not  depart  markedly  from  the  Pseitdacns  type. 

Feb.  2$,  1934.  Left  Kmston,  N.C.,  southward  on  No.  1 1  at  2:30  P.M.  Farther 
along  in  a  cattail  and  roadside  ditch  heard  a  few  isolated  Pscudacns.  Are  they 
Brandt's  P.  brimlcyi  or  what  ? 

Authorities9  corner: 
B.  B.  Brandt  and  C.  F.  Walker,  N.C.      B.  B.  Brandt,  N.C.,  1936^  p.  525. 

and  Ga.,  1933,  p.  5.  •  E.  B.  Chamberlain,  S.C.,  1939,  p.  17. 

B.  B.  Brandt,  N.C.,  19363,  p.  218. 

Swamp  Cricket  Frog,  Swamp  Chorus  Frog,  Swamp  Tree  Frog,  Swamp 

Tree  Toad,  Striped  Tree  Frog,  Rough  Chorus  Frog,  Le  Conte's 

Chorus  Frog,  Black  Chorus  Frog 

Pscudacns  nigrita  nigrita  (Le  Conte).  Plate  XLIX;  Map  17. 

Range:  North  Carolina  to  northern  Florida.  Mississippi,  possibly  into  Loui- 
siana. The  western  limits  are  not  very  clearly  defined. 

Habitat:  Breeds  in  ditches,  ponds,  or  bayous,  and  later  moves  to  dryer  ham- 
mocks and  ridges  of  the  pine  barrens. 

Size:  Adults,  %-iH  inches.  Males,  21-28  mm.  Females,  22-30  mm. 

General  appearance:  This  small  frog  has  a  slender  body  and  pointed  snout, 
prominent  eyes,  and  long  legs.  The  skin  is  finely  granulated.  This  form  has  the 
most  numerous  small  dorsal  spots  of  any  Psettdacns.  It  is  gray  or  olive  to  black 
in  color,  with  three  irregular  rows  of  dark  spots  on  the  back  and  darkly  mot- 
tled sides.  The  dark  spots  are  outlined  with  white  dots.  The  row  of  dark  spots 
down  mid-back  divides  into  two  rows  toward  the  rear.  A  light  line  is  present 
along  the  jaw  with  a  dark  area  extending  from  the  snout  through  the  eye  and 
beyond  the  tympanum.  There  is  a  light  speck  back  of  the  tympanum.  The  legs 
are  barred  with  black  spots.  There  is  no  triangle  or  crossbar  between  the  eyes. 
The  male  has  a  dark  greenish  yellow  throat  with  longitudinal  folds  in  the 
middle. 

Color:  Female.  Okefinokee  Swamp,  Ga.,  June  15,  1922.  Back  dark  gray, 
smoke  gray,  drab,  or  grayish  olive.  From  snout  down  middle  of  back  is  one 


TREE  FROGS:  HYLIDAE 


237 


238  HANDBOOK  OF  FROGS  AND  TOADS 

row  of  more  or  less  connected  dusky  drab  or  deep  brownish  drab  to  black 
spots.  Stripe  along  jaw  pale  rose  purple,  pale  salmon  color,  or  cartridge  buff 
with  some  seafoam  green. 

The  row  of  spots  continuous  down  the  back  becomes  at  halfway  point  two 
rows  of  separate  spots.  On  each  side  is  one  more  row  of  separate  spots.  Dark 
area  from  snout  through  eye,  back  of  eye,  and  extending  along  the  side  to 
groin.  Crossbars  of  legs  are  same  color  as  spots  of  back,  and  light  thin  bars  be- 
tween the  spots  of  legs  are  like  the  background  of  the  back.  Under  parts  white. 
Iris  black  with  brownish  vinaceous  dots.  At  times,  iris  has  considerable  glass 
green.  (See  A.  H.  Wright,  Gen.,  1932,  pp.  199-200  for  various  color  character- 
izations.) Sometimes  the  frogs  are  blackish  olive,  and  white  or  creamy  below. 
The  maxillary  stripe  may  be  white,  greenish,  or  a  little  yellow. 

Structure:  Slight  webs  at  bases  of  second,  third,  and  fourth  toes;  slight  disks 
at  tips  of  fingers  and  toes;  hind  legs  long;  in  many  ways  this  is  the  most  dis- 
tinctive of  all  six  subspecies  of  P.  mgnta. 

The  original  description — "Above  black  speckled  with  small  white  warts; 
middle  of  back  cinereous  with  an  interrupted  stripe  of  black,  upper  lips  with 
a  white  line;  beneath  granulate  whitish;  irides  golden;  legs  barred  with  whit- 
ish, hind  part  of  thighs  brown;  hind  legs  very  long"  (J.  Le  Conte,  Gen.,  1825, 

P.  282). 

Voice:  "Its  call  is  similar  to  that  of  the  Cricket  'frog,'  but  much  louder,  and 
the  crepitations  arc  slower"  (R.  F.  Deckcrt,  Fla.,  1915,  p.  23). 

In  1921  at  Fargo,  Ga.,  F.  Harper  noted  in  his  journal:  "During  downpours 
on  the  afternoon  of  August  5  ...  began  to  hear  several  frogs  calling  in  a  field 
grown  with  grasses  and  sedges  and  flooded  with  rain.  The  notes  consisted  of  a 
shrill  metallic  staccato  trill,  frequently  repeated:  i-i-i-ik,  i-i-i-ik,  i-i-i-ik.  They 
were  indistinguishable  from  those  of  representatives  of  this  genus  in  the  Atha- 
baska  region  (septentnonahs)  and  in  the  District  of  Columbia  (feriantm)" 

Breeding:  Breed  from  December  to  April.  Larval  period  40-60  days.  Trans- 
formation April  i-June  i  or  July.  Size  about  9-15  mm. 

Several  naturalists  in  the  South  have  made  notes  on  the  breeding  of  this  early 
species.  At  Biloxi,  M.  J.  Allen  (Miss.,  1932,  p.  7)  wrote:  "Several  pairs  were 
taken  in  copulation  on  Dec.  15,  1929  and  forty-two  breeding  pairs  and  some 
clumps  of  eggs  in  daylight  on  January  20, 1931." 

Carr  recorded:  "Breeding:  Jan.  16  to  April  2,  in  temporary  flatwoods  ponds 
and  ditches  and  in  the  shallow  margins  of  cypress  ponds.  January  and  Febru- 
ary choruses  almost  always  in  company  with  those  of  P.  ornata.  The  young 
emerge  in  April  and  May.  They  often  remain  for  several  weeks  about  the 
ponds  from  which  they  emerged"  (A.  F.  Carr,  Jr.,  Fla.,  1940,  p.  56). 

Journal  notes:  June  15, 1922,  Folkston,  Ga.  Near  Chesser  Schoolhouse  under 
chips,  small  boards,  and  logs  found  three  Psetidacns  in  dry  pine  barrens,  as- 
sociated with  Bufo  qtiercicus.  Area  beneath  boards  and  cover  more  or  less  dry. 


TREE  FROGS:  HYLIDAE 


Plate  XLJX.  PseudacrU  nigrita  nigrita 
XO.  i-4«  Adults. 


Plate  L.  Pseudacns  ntgnta  dar\ii  (XO- 
1,3,7.  Females.  2,6.  Males.  4.  Eggs  attached 
to  small  twig.  5.  Tadpole. 


246  HANDBOOK  OF  FROGS  AND  TOADS 

July  15,  Folkston,  Ga.  On  high  sandy  ridge  south  of  Spanish  Creek,  Quercus 
catesbaei,  Quercus  cinerea,  Gaylussacia  dumosa  and  a  very  much  serrated  poi- 
son ivy  (Rhtts)  were  growing.  Miles  and  I  were  looking  at  the  Rhus  carefully 
and  gingerly.  He  saw  something  jump  among  it.  Thought  it  a  grasshopper  but 
he  soon  found  it  to  be  a  frog.  It  was  Pseudacris  n.  nigrita.  We  took  its  picture 
right  there.  Other  plants  there  were  persimmon,  Baptisia,  brake,  and  croton. 
The  frog  was  first  on  a  dead  twig  of  a  fallen  branch.  It  would  leap  among  dead 
leaves,  among  wire  grasses,  and  among  the  ivy. 

Aug.  16.  In  making  a  detour  on  the  Dixie  Highway  about  a  mile  south  of 
Hilliard,  Fla.,  we  came  to  the  road  opposite  a  wet  pine  woods.  In  one  little 
temporary  pond  in  a  tussock  of  grass  beside  the  small  pool  was  a  Pseudacris 
n.  nigrita  calling.  To  me  this  form  which  was  finally  captured  sounded  like 
our  New  York  Pseudacris.  To  F.  Harper  it  sounded  like  a  metallic  staccato 
trill,  five  notes,  ic,  ic,  ic,  ic,  ic.  All  our  records  in  July  and  August  are  clearly 
sporadic  croakings  long  after  breeding  and  only  after  heavy  rainfall. 

Authorities9  corner: 

R.  F.  Deckert,  Fla.,  1915,  p.  23.    E.  B.  Chamberlain,  S.C.,  1939,  p.  15. 
C.  S.  Brimley,  N.C.,  1940,  p.  18. 

Clarke's  Chorus  Frog,  Clarke's  Striped  Tree  Frog,  Striped  Tree  Frog, 

Striped  Chorus  Frog 

Pseudacris  nigrita  clarion  (Baird).  Plate  L;  Map  18. 

Range:  Texas  to  Kansas.  "The  extension  of  the  range  of  the  species  north- 
ward has  been  a  gradual  and  a  continuous  one,  with  the  proper  interpretation 
of  the  recorded  species,  so  that  the  discovery  of  the  species  in  Kansas  is  not  so 
surprising.  Cope  extended  its  range  to  northern  Texas  (Dallas)  in  1880,  and 
into  Oklahoma  in  1894,  shortly  after  recording  it  from  the  panhandle  of  Texas 
(1893).  Ortenburger  then  (1926)  reported  nigrita  (almost  certainly  clar\ii,  by 
this  interpretation)  from  Creek  county,  Oklahoma,  a  record  verified  by  a  speci- 
men from  the  same  locality  in  the  K.  U.  Museum,  collected  by  Dr.  Taylor. 
Burt  (1932)  found  it  in  Kay  county,  Oklahoma,  and  subsequent  collecting 
and  examination  of  the  K.  U.  material  has  revealed  its  existence  far  northward 
into  central  Kansas,  in  essentially  the  same  ecological  relations  as  exist  where 
it  is  found  in  central  Oklahoma"  (H.  M.  Smith,  Kans.,  1934,  p.  465). 

Habitat:  Abundant  in  vicinity  of  marshes.  Breeds  in  roadside  ditches,  shal- 
low water-lily  ponds,  shallow  mesquite  ponds,  grassy  ponds,  or  other  transient 
pools. 

"Chorophilus  tn*eriatits  Wied.  .  .  .  Baird's  types  of  Helocoetes  clarion  were 
from  Indianola.  In  East-Central  Texas,  during  the  breeding  season,  this  species 
fairly  swarms  in  the  roadside  ditches  and  in  shallow  pools  on  the  grassy  flats" 
(J.  K.  Strecker,  Jr.,  Tex.,  1915,  p.  48). 


TREE  FROGS:  HYL1DAE  241 

"Chorophdtts  trisenatus  clarion.  Baird  and  Girard.  ...  An  abundant  spe- 
cies, especially  so  in  the  vicinity  of  marshes"  (J.  K.  Strecker,  Jr.,  Tex.,  1902, 

P-7)- 

"Abundant  and  noisy  in  pools  near  the  Cimmaron  River,  at  Tucker,  Okla- 
homa. Similar  to  the  individuals  reported  by  me  from  Clarendon,  Texas" 
(E.  D.  Cope,  Okla.,  1894,  p.  386). 

"The  specimens  found  in  Oklahoma  were  in  wooded  areas.  They  seemed  to 
avoid  the  deeper,  muddier,  less  protected  pasture  pools  where  Scaphiopus  and 
Bufo  were  numerous.  Some  were  found,  however,  hopping  about  m  the  grassy 
fields,  not  far  distant  from  pools.  The  specimens  from  Sedgwick  county,  Kan- 
sas, were  found  in  a  habitat  essentially  similar  to  the  above,  except  that  the  pool 
in  which  they  were  found  was  permanent,  while  the  other  apparently  was  not" 
(H.  M.  Smith,  Kans.,  1934,  p.  464), 

Size:  Adults,  %-i!4  inches.  Males,  20-29  mm-  Females,  25-51  mm. 

General  appearance:  This  is  a  small  grayish  or  olive  frog  with  dark  longi- 
tudinal spots,  or  these  spots  may  be  arranged  m  three  stripes.  A  dark  lateral 
banding  extends  to  the  nostril.  There  is  a  light  stripe  on  the  upper  jaw.  The 
legs  are  barred  above  and  white  or  pale  buff  beneath.  Venters  arc  white  or 
ivory  yellow  with  the  throat  of  the  male  a  dark  olive  buff.  They  are  protectively 
colored  little  things  in  the  grassy  edges  of  the  ditches  where  they  call  and  breed. 

Color:  Four  males.  Brownsville,  Tex.,  Feb.,  ig2$.  Upper  parts  light  grayish 
olive,  pale  olive-gray,  grayish  olive,  or  tea  green.  Spots  on  back  and  triangular 
spot  and  bars  on  legs  yellowish  oil  green,  oil  green,  or  courge  green.  Bar  from 
snout  through  nostril,  through  eye  and  tympanum,  and  onto  sides  black. 
Sometimes  back  of  shoulder  it  has  the  greenish  color  of  dorsal  spots.  Triangu- 
lar area  between  eyes  equilateral  in  one,  long-caudally  prolonged  in  another, 
divided  into  three  spots,  the  two  forward  angles  each  making  a  spot  and  the 
caudal  angle  prolonged  backward.  Area  in  another  not  a  triangle  but  with  pro- 
longation backward  at  one  end  like  a  figure  seven.  Two  of  these  four  have  a 
spot  ahead  of  the  triangle.  Above  lateral  vitta,  through  eye,  over  shoulder  is  a 
parallel  series  of  dorsal  spots,  usually  two,  the  caudal  one  longer,  or  the  cephalic 
one  larger,  or  all  one  or  occasionally  more  than  two.  If  there  are  other  dorsal 
spots,  they  are  in  the  posterior  half  of  the  back.  There  is  no  distinct  middorsal 
line  of  spots  or  line.  Belly  and  under  parts  white  except  for  throat,  which  is 
olive-ocher,  orange-citrine,  olive-yellow,  or  lemon  chrome. 

Detailed  color  descriptions  of  a  pair  taken  at  Bceville,  Tex.,  March  26,  1925, 
follow: 

Female.  Back  deep  olive-gray;  on  sides  pale  olive-gray  above  lateral  stripe. 
Spots  on  back  hellebore  green  surrounded  by  black.  Stripe  through  eye  and 
along  side  dark  grayish  olive.  Bars  and  spots  on  fore  limb  like  lateral  stripe. 
Stripe  on  upper  jaw  cartridge  buff  with  marguerite  yellow  or  primrose  yellow 
in  it.  Tympanum  brownish  olive  or  light  brownish  olive.  Venter  white,  pale 


242 


HANDBOOK  OF  FROGS  AND  TOADS 


olive-buff,  or  ivory  yellow.  Underside  of  hind  legs  light  grayish  vinaceous  or 
light  brownish  drab.  Iris— upper  part  of  rim  marguerite  yellow  or  primrose 
yellow,  rest  dotted  black,  primrose  yellow,  and  cinnamon  drab. 

Male.  Backgiound  pale  smoke  gray,  also  areas  of  tea  green.  Spots  vetiver 
green,  surrounded  by  black.  Lateral  stripe  bone  brown.  Tympanum  army 
brown.  White  or  primrose  yellow  spots  on  tubercles  below  vent  and  on  basal 
insertion  of  thighs.  Throat  discolored  dark  olive-buff  or  more  yellowish.  Two 
longitudinal  plaits  in  middle  of  throat.  Iris  of  this  male  has  also  some  avella- 
neous  in  it  and  a  tendency  to  a  dark  horizontal  bar  through  it  to  complete  nasal 
and  lateral  stripe. 

March  25,  1925.  In  the  two  pairs  which  laid,  the  females  were  more  spotted, 
the  males  more  striped.  Is  this  a  natural  tendency  or  an  individual  variation? 
The  male  of  a  third  pair  is  veiy  much  striped.  Another  male  we  captured  is 
very  much  spotted. 

Structure:  Snout  acute,  projecting  beyond  lower  jaw;  toes  slightly  dilated  at 
tip^;  male  throat  with  one  to  three  longitudinal  folds.  The  hind  limb,  tibia, 
snout,  and  head  are  longer  than  in  P.  n.  tnsenata. 

The  original  description — "Helocoetes  darken  Baird.  Snout  acute,  projecting. 
Extremities  somewhat  dilated.  Tibia  half  the  distance  between  eye  and  anus. 
Foot  but  little  longer,  not  nearly  half  the  length  of  body.  Above  grayish  brown 
or  ash,  with  distinct  large  circular  blotches.  A  dark  band  from  snout  through 
eye  and  tympanum  down  the  sides,  and  a  whitish  line  on  the  side  of  jaw.  Size 
about  one  inch  long.  Hub.  Galveston  and  Indianola,  Texas"  (S.  F.  Baird,  Gen., 
1854,  p.  *j). 

Voice:  The  call  is  a  grinding  note,  more  measured,  and  not  so  shrill  as  the 
call  of  Acns.  It  is  raucous  and  grating,  it-it  I  it-it /it-it,  sometimes  60  times  with- 
out a  stop,  sometimes  uniform,  sometimes  double  speed.  They  call  from  under 
the  grass  edges.  Sometimes  there  is  a  synchronized  chorus  of  sawing  notes. 

March  24,  1925,  Beeville,  Tex.  Air  is  resounding  with  Pseitdacrts.  8:30  P.M., 
air  74°.  Croaked  24  times,  stopped  instant,  then  began  again  25  to  30  times, 
hardly  catch  a  breath  then  start  again,  another  29  in  30  seconds.  They  are  hid- 
den under  the  grass  edges  like  Mtcrohyla,  which  is  calling  at  the  same  time  in 
the  same  roadside  ditches.  It  is  the  dominating  note  over  Mtcrohyla  and  Bufo 
valliceps  just  now. 

"Sometimes  in  later  afternoon  or  early  dusk,  preceding  a  heavy  chorus  dur- 
ing the  night,  P.  clarion  males  call  as  they  approach  a  breeding  pool  after  the 
manner  of  Hyla  v.  vemcohr.  These  calls,  however,  are  enough  different  from 
the  typical  breeding  cry  that  when  I  first  heard  them  I  was  not  sure  of  the  spe- 
cies responsible  till  I  had  run  down  and  captured  a  specimen.  Such  calls  are 
not  so  loud  nor  so  frequent  as  the  typical  breeding  call.  They  have  an  indescrib- 
able quality  which  for  want  of  a  better  term,  I  call  'pensive'  or  'pleading.'  Be- 
cause of  these  differences  I  suspect  (but  have  not  proved)  that  this  pensive  call 


TREE  FROGS:  HYIJDAE  243 

functions  to  attract  other  males  to  appropriate  breeding  pools  that  have  been 
sensed  by  males  nearer  to  them.  At  least,  such  calls  in  my  experience  have  been 
limited  to  a  circle  of  some  four  hundred  feet  in  diameter  whose  center  is  cov- 
ered by  water  appropriate  for  breeding  activities.  How  the  first  males  find  a 
pool  in  this  (and  other)  species  is  almost  entirely  unknown"  (A.  N.  Bragg, 
Okla.,  1943^  pp.  131-132). 

Breeding:  The  species  breeds  with  the  spring  rams,  March  5~May  20,  occa- 
sionally in  summer  to  mid-August.  The  citrine  drab  and  ivory  yellow  eggs  are 
in  a  loose  irregular  mass  attached  to  plant  stems,  their  number  few,  150  to  175 
eggs  in  groups  of  3  to  50.  The  egg  is  %2-/£r>  inch  (0.65-0.9  mm.),  the  outer 
envelope  M  2-M  n  inch  (2.2-2.4  mm-)  or  more  loose  and  irregular,  the  inner 
envelope  K«  inch  (1.4-1.8  mm.).  The  grayish  olive  tadpole  is  small,  TH-I/^ 
inch  (25-30  mm.),  its  crests  nearly  transparent,  the  tooth  ridges  %.  After  30  to 
45  days,  the  tadpoles  transform,  April  i  to  June  20,  at  r'i<j-/{>  inch  (cS-i  3  mm.). 
H.  M.  Smith's  (Kans.,  1934,  P-  4^4)  specimens  were  slightly  larger,  13.5-17.0 
mm.  in  length. 

One  pair  in  captivity  laid  154  eggs  in  several  small  masses  arranged  about 
one  inch  to  one  and  a  half  inches  apart  on  the  stem.  These  masses  were  17,  27, 
30,  37,  16,  13,  and  14  eggs,  respectively.  The  last  mass  rested  on  the  debris  in 
the  bottom  of  the  jar.  There  is  a  very  loose  outer  envelope,  at  times  irregular. 

We  found  the  tadpoles  transforming  in  extreme  southern  Texas  on  April  22. 
From  the  region  of  San  Antonio  we  have  the  following  note:  May  29,  1925. 
Some  Psetidacris  are  approaching  the  fully  developed  two-legged  stage.  Si'/c, 
8-13  mm. 

"Found  breeding  in  temporary  pools  from  early  May  to  mid-June.  Com- 
mon" (G.  A.  Moore  and  C.  C.  Rigney,  Okla.,  1942  [1941 1,  p.  78). 

"Feb.  25,  1944.  Heard  Pseiulacns  mgnta  clarlyi  frogs  at  Duck  Pond  about 
2^/2  miles  north  of  my  station.  These  arc  first  near  here.  Formerly  nearest  were 
near  Lytle  (9  miles)"  (A.  J.  Kirn,  Tex.). 

Journal  notes:  March  25,  1925,  Beeville,  Tex.  The  pair  we  caught  last  night 
broke  and  one  was  lost.  Captured  an  extra  female.  Put  in  photographic  jar,  two 
females  and  one  male.  This  morning  when  I  arose,  they  were  mated,  but  had 
not  laid.  The  female  of  this  pair  was  not  the  female  of  the  original  pair  cap- 
tured. They  laid  in  one-half  hour  or  less.  The  first  pair  captured  had  been  rest- 
ing at  the  grassy  edge  of  a  roadside  ditch.  The  female  was  larger  than  the  male. 
The  embrace  was  axillary. 

April  7, 1925.  The  eggs  laid  in  Bccville,  March  25,  are  developing  nicely. 

April  22,  San  Ben  i  to,  Tex.  In  sedges  around  a  pond  one  mile  south  of  San 
Benito,  these  little  transforming  frogs  were  swimming  in  shallow  water.  They 
often  duck.  Hard  to  collect  without  killing  them,  they  are  such  delicate  little 
creatures.  Skinned  hind  legs  of  two  in  their  capture.  They  can  turn  heads  like 
some  other  Psettdacris.  Found  only  a  very  few  tadpoles  in  the  pond.  The  species 


244  HANDBOOK  OF  FROGS  AND  TOADS 

must  be  through  metamorphosis  in  this  pond.  This  is  a  beautiful  water-lily 
pond  with  Marsilca  and  Castaha  elegans. 

Authorities'  corner: 
E.  D.  Cope,  Tex.,  1893,  P-  333- 
J.  K.  Strecker,  Jr.,  Tex.,  ipioc,  p.  80. 
A.  N.  Bragg,  Okla.,  1941,  p.  52. 

"The  comparison  of  these  two  tadpoles  \triseriata  and  darfyi]  reveals  an  in- 
teresting situation.  They  arc  so  nearly  alike  that  those  who  believe  them  of 
separate  species  (e.g.,  Smith  1934;  Bragg  1943)  can  take  little  comfort  from  the 
data.  However,  those  who  call  them  subspecies  or  varieties  of  one  species  (Burt, 
1932;  Wright  and  Wright,  1935, 1943;  Stejneger  and  Barbour,  1939)  must  ex' 
plain  the  proportional  differences  reported.  It  would  seem,  therefore,  that  either 

(1)  one  of  these  forms  has  fairly  recently  been  derived  from  the  other,  or 

(2)  both  have  been  derived  in  recent  geological  times  from  a  single  ancestral 
form.  The  latter  seems  the  more  probable  in  view  of  lack  of  intergradmg  speci- 
mens where  their  ranges  meet"  (A.  N.  Bragg,  Okla.,  I943C,  p.  138). 

Having  seen  most  of  the  tadpoles  of  the  United  States,  even  though  we  have 
not  described  them  all  minutely,  we  smile  at  "take  little  comfort  from  the 
data."  We  would  have  faith  today  in  no  one  who  pretended  he  could  distin- 
guish the  tadpoles  of  all  Psettdacris  or  all  Bttfo  species.  Bragg,  than  whom  no 
one  has  been  more  active  recently  in  life  histories,  must  have  enjoyed  writing 
the  paragraph  above.  P.  n.  clarfyi  and  P.  n.  triseriata  are  close.  When  we  find 
several  times  in  the  field  (not  in  alcohol)  pairs  of  P.  n.  clarfyi  with  the  males 
striped  and  the  females  spotted,  or  when  we  find  males  longer-legged  and  fe- 
males shorter-legged  in  some  P.  n.  septentnonahs  specimens,  it  means  that 
herpetologists  do  not  know  the  species;  or  original  descriptions  alone  do  not 
solve;  or  that  the  Pseudacris  nignta  complex  is  difficult  to  explain  by  any  classi- 
ficatory  schema  conceived  long  ago. 

Eastern  Chorus  Frog,  Eastern  Swamp  Cricket  Frog,  Swamp  Tree  Toad, 
Swamp  Tree  Frog,  Chorus  Frog,  Striped  Tree  Frog,  Common 

Chorus  Frog 

Pseudacris  nignta  jenartim  (Baird).  Plate  LI;  Map  17. 

Range:  New  Jersey,  Pennsylvania  to  North  Carolina,  and  possibly  into 
South  Carolina. 

Habitat:  Breed  in  marshy  stretches  or  shallow  ponds,  or  at  edges  of  ponds 
where  there  is  matted  vegetation,  sedgy  clumps,  or  moss. 

"Between  the  railroad  station  at  Annadale  and  Seguine's  pond  [Staten  Is- 
land j,  there  are  several  marshy  places  and  one  little  artificial  pond.  For  several 
years  we  have  heard  in  these  lowlands,  from  the  middle  of  March  to  the  first 
week  in  May,  the  song  of  what  we  considered  to  be  the  swamp  tree  frog,  Cho- 
rophilus  triseriattts  (Wied)"  (W.  T.  Davis,  N.Y.,  1912,  p.  66). 


TREE  FROGS:  HYLIDAE  245 

"This  species  I  have  found  abundant  on  the  sides  of  pools  and  ponds  in  the 
neighborhood  of  Gloucester,  N.J.,  in  the  spring  and  early  part  of  summer.  It 
delights  in  those  small  and  often  temporary  pieces  of  water  which  are  inclosed 
in  the  densest  thickets  of  spiny  Smilax  and  Rttbtts,  with  scrub  oaks,  and  sur- 
rounded by  the  water  loving  Cephalanthus,  where  no  shade  interrupts  the  full 
glow  of  sunlight"  (E.  D.  Cope,  Gen.,  1889,  p.  344). 

"As  the  pools  in  which  they  stay  are  usually  full  of  grass,  low  rushes  and 
other  vegetation,  they  are  not  easy  to  catch  although  patience  and  perseverance 
will  secure  specimens  without  too  much  trouble.  .  .  .  Outside  the  breeding 
season  these  frogs  are  not  often  seen,  but  may  be  sometimes  found  hopping 
about  in  damp  places  or  in  cool,  shady  woods.  On  the  ground  they  are  not 
much  more  active  than  toads"  (C.  S.  Bnmley,  N.C.,  1940,  p.  18). 

Size:  Adults,  %-i%  inches.  Males,  21-30  mm.  Females,  22-^  mm. 

General  appearance:  These  are  small  frogs  with  pointed  heads,  extremely 
long  fourth  toes,  very  little  or  no  webbing  between  the  toes,  and  with  small 
disks  on  their  toes.  They  are  grayish  olive  or  brown.  There  is  a  dark  triangle 
between  the  eyes  that  extends  backward  as  a  dorsal  stripe,  with  another  stripe 
on  either  side.  A  dark  stripe  extends  through  the  nostril  and  eye  to  the  shoul- 
der. This  is  bordered  below  with  a  light  stripe  on  the  upper  jaw.  The  throats 
of  the  males  are  usually  dark  but  not  always  so.  The  under  parts  are  a  buffy 
cream  color. 

Color:  Carlisle,  Pa.  Male.  Back  deep  grayish  olive,  drab,  or  drab-gray.  Tri- 
angle between  eyes  going  back  as  dorsal  stripe  with  a  stripe  on  either  side.  Each 
of  these  may  be  light-centered  with  body  color.  These  dorsal  stripes  may  be 
black,  grayish  olive,  light  grayish  olive,  or  deep  olive.  Undersides  of  hind  legs 
are  brownish  drab  or  benzo  brown.  Belly  cartridge  buff,  cream-buff,  or  cream 
color.  Throat  buckthorn  brown  near  tip  to  yellow  ocher  in  posterior  part.  Band 
from  snout  to  halfway  along  sides  mummy  brown  or  black.  Ins  same,  with 
little  burnt  sienna  in  upper  half.  Stripe  along  upper  jaw  almost  to  shoulder, 
warm  buff,  cream-buff,  or  chamois. 

Female.  Upper  part  of  head,  back,  forelegs,  and  hind  legs,  russet,  hazel, 
verona  brown,  or  warm  sepia.  Underside  of  hind  legs  benzo  brown,  drab-gray, 
or  brownish  drab.  Under  parts  of  body  cream  color,  becoming  slightly  massicot 
yellow  or  naphthalene  yellow  on  throat.  Stripe  down  middle  of  back,  stripe 
either  side  of  median  stripe,  and  faint  spots  on  sides  arc  black,  chestnut  brown, 
mummy  brown,  or  sepia.  Stripe  through  nostril  and  eye  to  shoulder  and  be- 
yond warm  sepia  or  Vandyke  brown.  Band  around  upper  jaw  extending  back- 
ward beyond  shoulder  below  the  dark  stripe  is  warm  buff,  cream  buff,  or 
chamois.  Iris  same  color  as  vitta  through  eye  but  dotted  with  mars  orange  or 
burnt  sienna.  These  predominate  in  the  upper  half,  giving  the  iris  a  "reddish" 
or  "coppery  red"  appearance. 

Structure:  The  original  description — "Helocoetes  feriarum,  Baird.  Body 
stout,  squat.  Head  broad.  Femur  and  tibia  and  hind  foot  about  equal,  and  half 


246  HANDBOOK  OF  FROGS  AND  TOADS 

the  length  of  the  body.  Above  dark  or  fawn,  with  three  nearly  parallel  stripes 
down  the  back,  the  central  widening,  but  scarcely  bifurcate  behind,  and  com- 
mencing behind  a  triangular  spot  between  the  eyes.  A  similar  dark  vitta  on 
sides  of  head  and  body,  with  a  white  line  along  edge  of  the  jaw.  Body  about  one 
inch  long.  Hab.  Carlisle,  Penna."  (S.  F.  Baird,  Gen.,  1854,  p.  60). 

Voice:  A  penetrating  and  grating  it-it/  it-it  is  given  many  times  without  a 
stop.  "Its  note  resembles  that  of  Acns  in  being  crcpitant,  and  differs  from  the 
toned  cry  or  whistle  of  the  Hylae.  It  is  not  so  loud  as  the  former  and  is  deeper 
pitched;  it  may  be  imitated  by  drawing  a  point  strongly  across  a  coarse  comb, 
commencing  at  the  bottom  of  a  jar  and  bringing  it  rapidly  to  the  mouth;  or 
better,  by  restraining  the  voice  to  the  separate  vibrations  of  the  vocal  cords,  and 
uttering  a  bar  of  a  dozen  or  twenty  vibrations,  beginning  with  the  mouth  closed 
and  ending  with  it  well  opened"  (E.  D.  Cope,  Gen.,  1889,  p.  345). 

"This  species  is  rarely  seen  except  during  its  breeding  season,  when  it  is  quite 
common.  Its  cry  consists  of  two  or  three  clear  whistlchke  chirps,  like  those  of 
a  young  turkey,  and  after  one  has  learned  to  distinguish  it  from  that  of  H.  pid^- 
eringi,  serves  to  betray  its  presence  at  once.  In  the  spring  of  1896  I  collected 
thirty  specimens  in  less  than  an  hour  from  the  gutters  along  the  Conduit  Road 
near  Cabin  John's  Bridge"  (W.  P.  Hay,  D.C.,  1902,  p.  129). 

"The  earliest  of  our  frogs  to  breed  at  Raleigh,  and  its  lazy  chorus  may  be 
heard  any  warm  day  in  February  and  March  and  even  in  December  and  Jan- 
uary, though  the  very  early  songsters  are  usually  only  single  individuals.  The 
cry  is  a  sort  of  grating  rattle  rising  at  the  end,  and  is  uttered  by  the  male  frogs 
lying  at  the  surface  of  some  shallow  pool.  They  are  very  shy  and  sensitive  and 
one  always  stops  singing  before  an  observer  gets  near  enough  to  perceive  the 
author  of  the  noise,  while  others  further  away  keep  up  the  chorus'*  (C.  S.  Brim- 
ley,  N.C.,  1940,  p.  T  8). 

April  15, 1921,  Raleigh,  N.C.  Temperature  of  air  68°F.  Tonight  at  9,  Vernon 
and  Julia  Haber  took  me  out  to  the  Oakwood  Cemetery  at  East  Raleigh  and 
to  St.  Augustine  Grounds.  Up  in  the  woodland,  where  we  couldn't  find  it,  was 
a  Pseudacns  fenantm  which  sounded  much  like  our  northern  swamp  cricket 
frog.  In  the  ponds  were  tadpoles  of  the  cricket  frog  in  all  stages  of  transforma; 
tion.  If  anything,  these  transform  at  a  little  larger  si/e  than  those  farther  north. 
In  order  of  activity  the  meadow  frog  and  Acns  gryllits  are  practically  through 
breeding,  though  in  the  ponds  the  swamp  crickets  have  left  for  uplands.  Fow- 
ler's toads  are  still  in  chorus  and  strongly  breeding  though  started  much  earlier, 
and  Hyla  versicolor  is  just  coming  to  the  pond. 

UI  have  heard  the  swamps  of  the  barrens  and  thickets  of  Southwestern  New 
Jersey  resound  with  them  as  early  as  the  twentieth  of  March,  when  a  skim  of 
ice  covered  part  of  the  water.  I  have  also  heard  it  in  other  level  parts  of  the 
same  State  later  in  the  season,  and  in  the  lower  part  of  Chester  County,  Pa." 
(E.D.  Cope,  1889,  p.  345). 

Breeding:  They  breed  from  February  to  May  10.  The  black  and  cream  eggs 


TREE  FROGS:  HYLIDAE  247 

arc  in  irregular  loose  jelly  masses  attached  to  the  stems  of  matted  vegetation. 
The  egg  is  %s  inch  (0.9-1.1  mm.),  the  single  envelope  %-/ii  inch  (3.2-4.0 
mm.).  The  vitelline  membrane  is  close  to  the  vitellus.  The  envelope  in  the  mass 
becomes  larger  and  irregular.  The  small  tadpole,  i-i'Kr*  inches  (25-33  mm-)» 
has  the  tail  medium,  with  tip  acute  or  acuminate,  and  is  blackish  or  olive  in 
color  with  bronzy  belly.  The  tooth  ridges  are  %.  After  50  to  60  days  the  tad- 
poles transform  until  mid-June,  at  Mc-Vs  inch  (8-12  mm.). 

April  6,  1929,  North  Mountain,  Carlisle,  Pa.  There  were  many  egg  masses 
attached  to  the  stems  in  this  vegetation  mat.  The  jelly  was  loose  and  indeter- 
minate in  shape.  It  stuck  to  the  stems  in  irregular  fashion  and  to  get  a  mass  a 
little  chunk  of  vegetation  had  to  be  pulled  out. 

"The  distribution  of  Pseudacns  is  limited  to  an  area  in  extreme  southern 
Berks  County  from  Gerger's  Mill  almost  to  Hopcwell  [N.J.]  and  south  to 
Elverson.  They  were  found  laying  eggs  at  Cold  Run,  May  4,  1937,  by  Earl  L. 
Poole.  Earliest  record:  calling,  Mar.  15,  1939.  Full  chorus  at  45°,  intermittent, 
scattered  calls  at  34°;  an  occasional  short  call  at  31°;  silent  at  ?o°FM  (C.  E. 
Mohr,  Pa.,  1939,  p.  78) . 

"The  eggs  arc  laid  in  small  masses  surrounded  by  a  very  weak  and  watery 
gelatinous  envelope  and  are  attached  to  the  stems  of  plants  or  other  supports 
just  below  the  surface  of  the  water  in  which  they  are  laid.  The  tadpoles  develop 
very  rapidly  and  I  have  found  them  as  early  as  April  16  with  the  hind  legs  suf- 
ficiently developed  for  them  to  be  able  to  hop  although  they  had  not  yet  lost 
their  tails"  (C.  S.  Brimlcy,  N.C.,  1940,  p.  18). 

Journal  notes:  April  3,  1929.  On  to  Carlisle,  through  Harnsburg.  Just  south 
of  Harrisburg  is  a  fine  grassy  shallow  pond  but  no  Pseudacns  were  calling 
there  by  day.  Drove  around  Carlisle  that  afternoon  to  spy  out  ponds  suitable 
for  Psetidacris.  We  heard  some  out  the  Mt.  Holly  Rd.  in  a  punched  tussock 
and  swampy  area  in  a  field.  Other  ponds  between  Mt.  Holly  and  Dillsburg 
had  many  peepers  but  no  Pseudacns.  Went  out  the  Chambershurg  Rd.  (April 
4)  and  on  the  way  back  found  a  surface  pond  at  the  edge  of  town,  in  fact  in 
a  vacant  lot  near  a  big  building.  Heard  just  three  Pseudacns  here  and  caught 
them  all.  We  hated  to  give  up  with  only  three  real  Carlisle  Pseudacns,  but  the 
next  night  brought  out  no  more. 

April  6,  started  up  North  Mountain.  Just  before  Indian  Springs,  well  up  on 
the  mountain,  we  heard  a  great  chorus  of  Psetidacris  feriarum.  On  the  hillside, 
beside  the  road  in  an  open  field  was  a  springy  area.  Finally  we  saw  two  little 
fellows  with  their  amber  throats  so  distended  that  they  showed  from  the  rear. 
They  were  sitting  in  the  mat  of  vegetation,  a  little  ways  from  a  tussock,  with 
their  heads  held  up  vertically,  above  the  water.  We  went  down  to  catch  them 
and  found  the  stream  area  fairly  deep  but  filled  with  this  mat  of  grassy  vegeta- 
tion. We  rolled  this  back  a  bit  at  a  time  and  caught  an  occasional  frog  in  the 
tangle.  One  was  a  female,  ripe,  and  strange  to  say  she  looked  slender,  as  did 
the  males  from  the  Carlisle  Pond,  while  the  males  from  this  Mountain  Spring 


248  HANDBOOK  OF  FROGS  AND  TOADS 

area  were  fat  and  squatty.  We  caught  six  or  seven  males  and  one  female  there. 

June  10, 1929.  Over  the  hills  from  Scranton  to  Harrisburg,  leaving  the  main 
road  at  Clark's  Ferry,  and  over  the  mountain  toward  Carlisle,  back  to  the  spot, 
Indian  Springs  Garden,  where  we  caught  Psetidacris  n.  feriarum  at  Easter 
time.  Our  springy  hillside  is  almost  dried  up  but  is  still  spongy  and  water  stands 
where  you  step,  but  no  Pseudacris  visible.  Looked  for  little  transformed  ones 
but  found  none.  Bert  has  gone  down  to  the  pond  in  the  meadow  below;  yellow 
sweet  clover,  red  clover,  daisies,  wild  roses  are  in  bloom.  No  luck  at  all.  On 
toward  Carlisle  with  crestfallen  faces.  A  pond  by  the  roadside— many  tadpoles, 
but  not  the  right  ones.  Came  to  a  small  stream  with  culvert  in  the  road,  a  deep- 
cut  stream  with  6-foot  banks.  Above  one  bank  was  a  springy  inlet  with  masses 
of  sensitive  fern,  grassy  tussocks,  touch-me-not,  smartweed,  and  a  little  sedge. 
To  the  left  in  quiet  water  pockets  sometimes  no  more  than  4  to  6  inches  deep, 
among  grassy  tussocks  and  a  little  sedge  and  fern  was  our  big  game.  Tadpoles 
in  general  quite  dark.  As  they  approach  transformation  the  triangle  between 
the  eyes  becomes  quite  prominent.  It  curves  in  on  each  side,  and  the  rear  point 
extends  as  a  line  down  the  back  to  the  sacral  region.  On  either  side  of  middorsal 
stripe  appears  a  dorsolatcral  one  of  the  same  length.  As  the  tads  rest  in  the 
water,  the  tail  region  seems  to  be  more  or  less  speckled  or  to  have  an  alterna- 
tion of  light  and  dark.  We  were  impressed  with  long  tail  and  bronzy  belly. 
Some  tadpoles  were  only  half  grown. 

Authorities1  corner: 
R.  Sutcliff,  Gen.,  1812,  pp.  213-214. 
W.  T.  Davis,  N.Y.,  1912,  pp.  66-67. 

Queries  and  comments:  i.  Good  authorities  have  identified  specimens  from 
eastern  Tennessee,  Arkansas,  and  Illinois  as  P.  n.  feriarum,  and  Del-Mar-Va 
specimens  as  P.  n.  triseriata.  Are  P.  n.  feriarum  and  P.  n.  triseriata  one? 

2.  In  1921,  we  became  acquainted  with  P.  n.  feriarum  at  Raleigh,  N.C.,  and 
in  1934  we  saw  in  Washington,  N.C.,  live  tadpoles  and  material  of  P.  brimleyi. 
Much  remains  to  be  done  on  these  two  forms. 

$.  Our  Carlisle,  Pa.  (type  locality),  material  did  not  correspond  too  exactly 
with  Baird's  description. 

4.  For  30  to  40  years  our  students  from  Philadelphia  and  New  York  City 
have  from  time  to  time  queried  whether  there  were  two  different  forms  in 
New  Jersey. 

5.  Is  P.  n.  feriarum  coastal  in  the  northern  and  Piedmont  portions  and  inland 
in  the  southern  portion  of  its  range? 

6.  Does  P.  n.  feriarum  really  reach  northwestern  Florida?  The  Tampa  Bay 
record  must  be  a  mistake. 


TREE  FROGS:  HYLIDAE 


249 


Plate  LI.  Pseudacris  nigrita  feriarum  Plate  L/7.  Pseudacris  nigrita  septcntrh- 

(XO- 1,2,6-  Females.  3,5,7.  Males.  4.  Eggs.          nalis.  1,3,4.  Females  (Xi).  2.  Male  (XO- 

5.  Female  (Xi%). 


250  HANDBOOK  OF  FROGS  AND  TOADS 

Northern  Striped  Tree  Frog,  Northern  Chorophilus,  Northern  Pseudacris, 
Spotted  Tree  Frog,  Northern  Spring  Peeper,  Peeper  Frog,  Swamp 

Whistler 

Vscttdacris  nigrita  septentnonalis  (Boulcnger).  Plate  LII;  Map  18. 

Range:  Minnesota  to  Hudson  Bay,  to  upper  British  Columbia  and  Macken- 
zie through  Montana,  Saskatchewan,  and  Alberta;  Canadian  Northwest.  Ed- 
ward A.  Preble  found  it  from  Lake  Winnipeg  to  York  Factory  on  Hudson 
Bay  and  from  Edmonton,  Alberta,  to  Ft.  Norman  west  of  Great  Bear  Lake. 
Some  authors  and  collectors  have  felt  inclined  to  assign  some  of  the  swamp 
cricket  frogs  of  our  northwest  border  states  from  Minnesota  to  Montana  to 
this  form,  and  surely  some  of  the  specimens  we  collected  in  the  Red  River  of 
the  North  were  short-legged  enough  to  fall  withm  this  subspecies.  G.  C.  Carl 
(B.C.,  1945)  reports  it  for  Peace  River  district. 

Habitat:  Swampy  borders  of  rivers,  lakes,  ponds,  and  meadows. 

Size:  Adults,  %-i%  inches.  Males,  19-32  mm.  Females,  19-35  ram. 

General  appearance:  Pembma,  N.D.  This  is  a  small,  long-bodied,  short- 
legged  tree  frog.  The  snout  is  pointed.  The  color  may  vary  from  a  gray  to 
tawny  or  buff.  The  most  prominent  marks  are  the  dark  brown  lateral  stripe 
from  snout  to  groin,  and  the  light  creamy  stripe  on  the  jaw  below  the  eye.  Cer- 
tainly when  seen  at  rest,  you  are  impressed  with  the  very  short  hind  legs  in 
sharp  contrast  with  the  long  body.  It  has  five  darker  stripes  down  the  body. 
The  median  dorsal  one  may  be  broken  into  dots,  the  two  laterodorsal  stripes 
are  more  constant  and  the  lateral  ones  most  prominent.  The  male  has  a  green- 
ish yellow  throat,  the  female,  a  light  throat.  The  legs  of  the  female  arc  short 
and  stout,  those  of  the  male  longer  and  thinner.  The  under  parts  are  greenish 
white. 

Color:  "The  following  brief  description  of  the  color  was  taken  from  a  live 
specimen  from  Oxford  House:  Body  light  green  above,  greenish  white  be- 
neath; body  stripes  bronzy  lavender;  tympanum  brownish;  hind  legs  light 
green  above,  flesh  color  beneath*'  (E.  A.  Preble,  Kcew.,  1902,  p.  134). 

Females.  Pembina,  N.D.,  Aug.  30,  1930.  First  female.  The  color  of  top  of 
head,  top  of  arms  and  legs,  and  back  between  the  three  dorsal  stripes  is  drab*. 
The  color  between  outer  dorsal  stripe  and  lateral  band  is  a  smoke  gray  or  pale 
drab-gray.  The  band  extending  from  snout  through  eye  and  over  tympanum 
to  near  groin  is  olive-brown  or  brownish  olive.  This  color  is  solid  to  the  arm 
insertion,  then  goes  over  a  pebbly  surface,  the  interstices  of  which  are  smoke 
gray  or  pale  drab-gray,  like  the  color  above  the  band.  The  median  and  two 
laterodorsal  stripes  are  huffy  brown  outlined  with  dotted  lines  of  olive-brown. 
On  top  of  head  arc  a  few  specks  of  olive-brown.  The  femur  and  tibia  also  have 
spots  of  olive-brown,  and  on  front  of  tibia  and  rear  of  foot  and  tarsus  are  half 
bars  of  the  same  color.  On  the  upper  jaw  there  is  a  prominent  line  of  olive-buff 
or  chartreuse  yellow  bordered  below,  broadly  on  upper  jaw  and  narrowly  on 


TREE  FROGS:  HYLIDAE  251 

lower  jaw,  by  a  finely  punctate  band  of  buffy  olive  or  drab.  Under  surfaces  are 
pale  dull  green-yellow,  except  for  underside  of  hind  legs,  which  is  light  grayish 
vinaceous.  All  five  stripes  are  continuous.  The  eye  is  olive-brown,  except  for 
the  upper  rim  of  pupil,  which  is  clear  yellow-green;  the  ins  is  dotted  all  over 
with  clear  yellow-green  and  mars  orange  spots;  the  front,  back,  and  lower  mar- 
gin of  pupil  rim  is  a  succession  of  clear  yellow-green  dots. 

Second  female.  In  middle  of  back  the  dorsal  stripe  breaks  into  two  irregular 
rows  of  small  spots. 

Third  female.  The  back  is  tawny-olive  to  ochraceous-buff.  The  dorsal  stripes 
are  indistinct  but  nevertheless  indicated.  The  lateral  stripe  is  most  prominent. 

Male.  Walhalla,  N.D.,  Sept.  i,  1950.  The  area  between  lateral  band  and 
laterodorsal  stripe  is  pale  olivine  or  pale  fluonte  green.  The  top  of  back  is  mi- 
gnonette green.  The  three  dorsal  stripes  are  Kronberg's  green  surrounded  by 
broken  lines  of  olive-brown.  The  same  color  extends  from  snout  through  eye 
and  halfway  along  the  side.  The  stripe  below  the  eye  is  glass  green.  The  eye  is 
like  the  lateral  stripe  in  color  and  has  the  same  dots  as  the  female's,  except  that 
the  upper  iris  rim  is  primulme  yellow.  Under  parts  are  pale  niagara  green.  The 
throat  is  citrine,  old  gold,  or  olive  lake. 

Structure:  Hind  legs  short.  The  original  description — "Chorophtlus  septen- 
tnonalis.  .  .  .  Psendacns  mgnta,  part  T,  Gunth.  Cat.  p.  97.  Tongue  oval, 
slightly  nicked.  Vomerine  teeth  in  two  small  groups  behind  the  level  of  the 
choanae.  Head  longer  than  broad;  snout  subacummate,  prominent,  twice  as 
long  as  the  diameter  of  the  eye;  latter  very  small,  can  thus  rostrahs  rather  indis- 
tinct; interorbital  space  a  little  broader  than  the  upper  eyelid;  tympanum 
nearly  as  large  as  the  eye.  Fingers  and  toes  moderately  slender,  latter  with  a 
slight  rudiment  of  web;  first  finger  shorter  than  second;  an  indistinct  outer 
mctatarsal  tubercle.  The  hind  limb  being  carried  forwards  along  the  body,  the 
tibio-tarsal  articulation  reaches  hardly  the  tympanum.  Skin  granulate  above 
and  beneath.  Yellowish  olive  above,  with  five  longitudinal  dark  bands — the 
median  bifurcating  on  the  sacral  region,  the  outermost  extending  from  the  tip 
of  the  snout  through  the  eye  to  the  middle  of  the  side;  beneath  whitish,  im- 
maculate. From  snout  to  vent  25  milhm.  ...  a.  9  Great  Bear  Lake.  Well 
distinguished  from  C.  tnseriatus  by  its  shorter  hind  limbs"  (G.  A.  Boulcnger, 
Gen.,  1882,  pp.  335-556). 

Boulenger's  key  separates  this  form  from  mgnta  and  tnseriata  by  its  rough 
dorsal  region. 

Voice:  Its  call  is  like  those  of  the  more  southern  forms. 

"This  little  frog,  whose  trilling  notes  arc  exactly  similar  to  those  of  the  more 
southern  forms  of  the  genus,  is  fairly  common  nearly  throughout  the  region 
[Athabaska-Mackenzie  Region].  ...  At  Fort  Simpson  I  first  heard  its  notes 
on  May  3, 1904,  but  failed  to  take  specimens.  I  heard  it  also  on  the  Mackenzie 
above  Fort  Norman  early  in  June"  (E.  A.  Preble,  Athab.,  1908,  p.  502). 

"Though  its  piercing  'prreep  prreep!  from  the  chilly  pond,  in  early  spring- 


252  HANDBOOK  OF  FROGS  AND  TOADS 

time  is  familiar  to  all,  very  few  have  seen  the  originator  of  the  noise  or  know 
that  it  is  a  tiny  frog  that  makes  this  small  steam-whistle.  While  uttering  it,  his 
throat  is  blown  out  like  a  transparent  bladder,  and  is  nearly  as  big  as  himself. 
At  Shoal  Lake,  in  1901, 1  found  them  still  singing  in  the  first  week  of  July.  The 
note  is  more  rattled  than  that  of  H.  crucifer"  (E.  T.  Seton,  Man.,  1918,  p.  83) . 

"Northern  Frog.  Pseiulacris—igiq:  June  18,  heard  at  One  Four;  Milk  River 
July  21,  one  and  again  on  Aug.  4.  Common  about  McGrath  from  middle  to 
end  of  August"  (M.  Y.  Williams,  Alta.  and  Sask.,  1946,  p.  49). 

Harper  (  Athab.,  1931,  pp.  68-70)  heard  them  trilling  from  May  1 1  to  Oct.  21. 

Breeding:  This  species  breeds  from  May  to  early  June.  The  eggs  arc  in  an 
irregular  jelly  mass  attached  to  vegetation.  The  eggs  are  Viio-Mo  inch  (1.2-1.4 
mm.).  The  tadpoles  transform  at  about  %«-%  inch  (7.5-13  mm.). 

In  Manitoba  E.  T.  Seton  has  taken  several  immature  specimens  between 
14.5  and  17.0  mm.  R.  Kennicott  at  Ft.  Resolution,  Canada,  secured  immaturcs 
as  small  as  16.5-19.0  mm.  In  Minnesota,  in  the  valley  of  the  Red  River  of  the 
North,  the  same  collector  secured  immatures  15.0  and  15.5  mm.  Young  of  the 
latter  size  were  secured  by  J.  P.  Jensen,  September,  1908,  at  Eagle  Bend,  Min- 
nesota. In  Montana  at  the  Crow  Agency  M.  A.  Hanna  secured  a  similar  set 
16-18  mm.  on  July  i  T,  1916.  All  these  must  be  well  beyond  transformation  size, 
for  three  out  of  the  five  nigrita  subspecies  we  know  transform  at  7.5-13  mm. 

Journal  notes:  Aug.  31, 1930,  Red  River  of  the  North,  Pembma,  N.D.  Started 
out  about  8  A.M.  to  beat  the  vegetation  along  the  river  bank,  expecting  to  find 
more  northern  wood  frogs.  Along  the  shore  is  a  mud  flat,  baked  and  broken 
into  squares.  The  cracks  are  deep  and  fairly  wide.  There  are  deeper  holes 
punched  by  cows  and  horses.  Many  crickets  and  spiders  were  running  over 
the  flats,  and  clouds  of  small  insects  come  out  of  the  more  moist  footprints.  We 
walked  along  the  shore,  more  or  less  abreast;  Bert  was  in  the  vegetation  and 
I  walked  on  the  flats  at  the  edge  of  the  vegetation.  I  beat  the  bordering  vegeta- 
tion with  a  cane  and  searched  particularly  along  the  mud  at  the  edge,  expecting 
at  any  moment  to  sec  a  wood  frog.  Suddenly  my  eye  spied  a  small  frog  sitting 
at  the  edge  of  one  of  the  clumps.  As  I  looked  at  it,  it  jumped  into  a  crevice.  As 
I  looked  closely,  imagine  my  surprise  to  find  that  it  was  a  Pseudacns  n.  septen- 
trionalis.  We  have  been  looking  for  suitable  places  for  the  search  of  this  specie* 
for  some  days.  Soon  after,  I  saw  a  second  clinging  to  the  edge  of  another  crev- 
ice. As  we  came  close  to  catch  it,  it  cocked  its  head  to  one  side  as  some  other 
Pseudacris  do  at  times.  A  little  later,  a  third  appeared  clinging  to  a  close-grown 
grassy  edge  covering  clumps  bordering  the  mud  flats.  All  three  arc  females. 
They  certainly  have  short  hind  legs.  In  no  case  did  we  get  the  impression  of  the 
frog  leaping.  They  must  have  moved  to  have  caught  my  eye,  but  it  was  not  a 
leap  like  that  of  a  wood  frog  or  meadow  frog.  The  last  one  acted  as  if  crawling 
up  the  grass.  In  captivity  they  leap  from  2  to  8  inches.  There  are  a  few  meadow 
frogs  along  the  shore  and  we  caught  several  northern  wood  frogs. 


TREE  FROGS:  HYLIDAE  153 

Sept.  i,  1930,  Walhalla,  N.D.  The  boys  in  searching  the  manholes  of  drain- 
age sewers  saw  a  midget  on  the  side  wall  of  one  hole  and  with  searchlight 
climbed  down  the  ladder  after  it.  It  proved  to  be  the  much-desired  game,  a 
male  Pseudacris  n.  septentrionalts.  It  is  smaller  than  the  females  from  Pembina 
and  the  hind  legs  look  longer  and  slimmer. 

Authorities9  corner: 

L.  Agassiz,  Ont.,  1850,  p.  379.  G.  C.  Carl,  B.C.,  1943,  p.  46. 

E.  T.  Seton,  Man.,  1918,  pp.  82-83.  W.  J.  Breckenridge,  Minn.,  1944,  p.  67. 

I.  McT.  Cowan,  B.C.,  1939,  p.  92. 

Striped  Tree  Frog,  Three-striped  Tree  Frog,  Swamp  Cricket  Frog,  Striped 

Bush  Frog,  Three-lined  Tree  Frog,  Swamp  Tree  Frog,  Peeper,  Spring 

Peeper,  Western  Striped  Frog,  Western  Marsh  Toad 

Pseudacris  mgnta  tnsenata  (Wied).  Plate  LIII;  Maps  17,  18. 

Range:  Along  the  shores  of  Lake  Ontario  westward  through  Ontario, 
Michigan,  Wisconsin,  Minnesota,  North  Dakota,  Montana,  to  Idaho  south- 
ward to  northern  Arizona  and  northern  New  Mexico.  No  records  in  western 
Nebraska  and  Kansas.  Eastern  Kansas  southeastward  through  Oklahoma  to 
Arkansas  and  eastern  Texas,  possibly  to  northern  Louisiana,  thence  through 
Tennessee  and  Kentucky  (few  records)  to  Ohio  and  western  Pennsylvania. 

Habitat:  Low  bushes  and  plants,  and  on  the  ground.  Breeds  in  ditches, 
swamps,  or  temporary  ponds. 

"This  species,  though  called  a  tree-frog,  probably  never  climbs  trees.  They 
seem  to  live  on  the  ground  among  the  fallen  forest  leaves  and  in  the  grass" 
(O.P,Hay,Ind.,i892,p.47i). 

"This  is  one  of  the  nicest  marked  of  the  small  frogs,  found  generally  in  early 
spring  in  the  ditches  along  the  country  roads,  Wet  Prairie,  Madison  County, 
111.,  and  near  Bluff  Lake,  St.  Clair  County,  111.  It  is  often  seen  during  harvesting 
season  and  for  that  reason  farmers  call  it  the  'Hay  Frog'  "  (J.  Hurter,  111.,  1893, 
p.  254). 

"This  species  is  almost  always  found  on  the  ground,  where  it  hides  away 
under  loose  stones  or  fallen  timber"  (M.  M.  Ellis  and  J.  Henderson,  Colo.,  191 3, 

P.  SB). 

"It  is  to  be  found  in  swamps  on  low  herbage  or  on  the  ground"  (M.  Morse, 
Ohio,  1904,  p.  119). 

Size:  Adults,  %-iV«>  inches.  Males,  21-32  mm.  Females,  20.0-37.5  mm. 

General  appearance:  These  are  small,  slender  frogs  with  pointed  heads,  ex- 
tremely long  fourth  toes,  and  small  disks  on  their  toes.  They  are  brown,  olive, 
or  grayish  with  a  dark  brown  triangle,  spot,  or  stripe  between  the  eyes.  The 
most  prominent  stripe  is  one  from  the  nostril  through  the  eye,  over  the  arm, 
and  extending  along  the  side  halfway  to  the  groin.  There  are  three  dark  stripes 


HANDBOOK  OF  FROGS  AND  TOADS 

down  the  back,  continuous  or  more  or  less  broken  up,  and  dark  spots  on  the 
legs.  There  is  a  prominent  light  cream  or  silvery  line  along  the  upper  jaw.  They 
are  light  cream  or  white  beneath.  Their  skm  is  finely  granular. 

April,  1928,  Hilton,  N.Y.  At  night  it  looks  mud-colored.  Seldom  did  the 
stripes  or  spots  appear  much  revealed.  Pointed  head,  long  legs,  small  and  deli- 
cate in  form.  When  perched  on  a  stick  or  floating  mass,  sometimes  reminds 
me  of  P.  oculans. 

Color:  Ann  Arbor,  Mich.,  from  Mrs.  H.  T.  Gaige,  April  7-8, 1929.  Female. 
Sayal  brown,  tawny-olive,  or  wood  brown  on  back  and  upper  part  of  fore 
and  hind  legs  and  tip  of  head.  Stripes  and  spots  bone  brown,  black,  or  chest- 
nut brown.  The  marks  are:  a  triangle  at  the  eyes  extending  backward  as 
a  median  stripe,  which  may  be  interrupted  and  resumed  and  subdivided  on 
rear  back  into  two  parallel  lines;  another  band  along  dorsolateral  region  from 
above  shoulder  to  groin;  and  a  stripe  from  snout  through  nostril,  eye,  and  tym- 
panum halfway  to  groin.  Tympanum  snuff  brown  or  mikado  brown.  Stripe 
on  upper  jaw  warm  buff  or  cream-buff  with  upper  part  light  pinkish  cinnamon 
below  dark  stripe  through  nostril  and  eye.  Dark  edge  below  this  light  stripe 
is  on  labial  border.  Iris  same  color  as  stripe  along  head,  with  dottings  of  mars 
orange  or  burnt  sienna;  over  pupil  rim  is  light  yellow-green.  Under  parts  as  in 
P.  n.  jerarium  from  Carlisle,  Pa.,  are  cream  color,  becoming  slightly  massicot 
yellow  or  naphthaline  yellow  on  throat. 

Male.  Black  above  (only  under  lens  are  any  stripes  revealed,  except  stripe 
from  snout  through  eye  halfway  to  groin,  which  is  black).  Light  stripe  on 
upper  jaw  as  in  female.  Throat  yellow  ocher  or  old  gold.  Venter  white.  Under- 
side of  hind  legs  as  m  female.  While  in  hand  the  upper  parts  changed  to  buffy 
olive  or  grayish  olive. 

Structure:  Vocal  pouch  round  and  subgular. 

"On  the  forefoot  the  second  toe  (counting  from  the  outside)  is  much  longer 
than  the  others,  which  are  short;  on  the  hind  foot,  very  much  longer  still;  webs 
of  the  hind  feet  take  up  not  quite  half  the  room  between  the  toes;  ins  golden 
colored;  a  dark  brown  stripe  runs  from  the  nose  through  the  eye  up  to  the 
hind  leg;  the  edge  of  the  upper  jaw  white;  all  the  upper  part  an  inconspicuous, 
dark  greenish  gray,  with  three  longitudinal  rows  of  large,  elongated  spots  of 
the  same  hue  (only  somewhat  darker)  on  the  back,  .  .  .  belly  white;  throat 
and  chin  brownish-olive  colored;  under  side  of  the  hind  legs  reddish  gray.  .  .  . 
Length  il//i  length  with  hind  legs  extended  2'g";  length  of  the  longest  fore- 
toe  21/4";  length  of  the  longest  hind  toe  4%"  "  (M.  zu  Wied,  Gen.,  1839,  I, 
249-250). 

Voice:  The  call  is  a  vibrating  chirp. 

"During  the  last  weeks  of  March  Chorophilus  (Pseudacris)  appears  in  con- 
siderable numbers  about  the  outskirts  of  Buffalo.  The  male  chorus,  which  is 
easily  distinguished  from  that  of  Hyla  pic\enngii,  rises  from  most  of  the 
swamps  and  temporary  ponds,  even  within  the  city.  The  singers  themselves, 


TREE  FROGS:  HYLIDAE  255 

however,  are  not  easily  seen,  for,  upon  approach,  they  become  silent  and  further 
disturbance  causes  them  to  disappear  into  the  vegetation  at  the  bottom  of  the 
pond  where  they  remain  until  some  time  after  the  disturbance  has  ceased. 
Then,  from  some  remote  corner  the  chorus  is  gradually  taken  up  until  the 
whole  pond  resounds  with  the  ringing  notes.  In  taking  up  the  chorus,  the  as- 
surance evidenced  by  the  single  voice  is  extremely  contagious.  This  fact  makes 
it  possible  to  overcome  some  of  the  difficulties  which  ordinarily  present  them- 
selves in  collecting  this  species.  One  has  but  to  place  the  first  captures  in  a  bag 
or  other  close  receptacle  and  carry  it  on  one's  person.  The  prisoners  chirp  up 
and  sing  as  boldly  as  though  undisturbed  in  their  natural  haunt.  Their  voices 
elicit  an  almost  immediate  response  from  those  in  the  pond.  Indeed,  at  such  a 
time,  with  a  little  practice  one  can  wade  about,  while  they  sing  on  all  sides  and 
dip  up  as  many  as  one  desires.  In  this  way  after  spending  several  hours  in  cap- 
turing the  first  two,  30  specimens,  25  males  and  5  females,  were  taken  in  less 
than  an  hour"  (A.  H.  Wright  and  A.  A.  Allen,  N.Y.,  1908,  pp.  39-40). 

"It  has  a  note  somewhat  similar  to  the  preceding  species  (Acns  gryllus  crept- 
tans),  but  the  pitch  is  higher  and  the  rattle  is  less  definite.  The  note  is  seldom 
heard  in  daylight  hours  except  on  dark  days.  The  writer  has  never  heard  it, 
as  Cope  &ays,  in  the  hottest  hours"  (M.  Morse,  Ohio,  1904,  p.  119). 

"With  the  first  mild  spring  days,  often  before  all  the  snow  and  ice  of  winter 
have  disappeared,  the  loud  trill  of  this  small  species  may  be  heard  from  pools 
and  ditches.  The  note  is  so  resonant  that  on  quiet  evenings  it  may  be  heard  a 
half  mile  or  more  and  is  commonly  attributed  to  the  larger  frogs  of  the  genus 
Rana.  When  the  note  is  uttered  the  vocal  sac  is  extended  to  its  utmost  and  is 
larger  than  the  head"  (H.  Garman,  111.,  1892,  p.  345). 

"The  swamp  tree  frog  is  found  in  marshes  and  damp  places  throughout  the 
summer  and  fall.  During  this  time  it  is  solitary  and  its  call  is  rarely  heard.  It  is 
also  seldom  seen  because  of  its  small  M/C  and  protective  coloration.  When  dis- 
turbed it  disappears  in  the  water,  but  it  is  a  very  poor  swimmer  and  soon  comes 
back  to  land.  ...  It  comes  from  hibernation  early.  The  song  is  very  loud. 
When  croaking,  the  male  sits  upright  in  the  water,  supporting  himself  with 
grass,  leaves  or  twigs,  and  sings  with  the  head  and  vocal  pouch  out  of  the  water. 
When  disturbed,  he  sits  perfectly  still  and  does  not  resume  his  song  until 
the  source  of  the  alarm  has  passed"  (A.  G.  Ruthven,  C.  Thompson,  and  H. 
Thompson,  Mich.,  1912,  pp.  47-48). 

Breeding:  This  species  breeds  from  March  20  to  May  20.  The  egg  mass  is  a 
loose  irregular  cluster,  the  mass  small,  less  than  i  inch  in  diameter,  20-100  eggs 
in  the  mass,  500-1500  eggs  in  the  complement.  The  brown  or  black  and  white 
egg  is  /^ri-%0  inch  (0.9-1.2  mm.),  the  single  envelope  M»-%o  inch  (5.0-7.8 
mm.).  The  tadpole  is  small,  i:Ko  inch  (23  mm.),  deep-bodied  with  a  long 
tipped  tail,  quite  black  with  bronze  on  the  belly  and  sides,  and  the  tooth  ridges 
are  %.  After  40  to  90  days,  the  tadpoles  transform  in  June  at  /io-%o  inch  (7.5- 
ii  mm.). 


256  HANDBOOK  OF  FROGS  AND  TOADS 

"While  travelling  along  the  state  highway  in  the  southern  part  of  Logan, 
Utah,  on  the  i5th  of  May,  1919,  I  heard  great  numbers  of  the  swamp  tree 
frog,  Pseudacris  triscriata,  uttering  their  characteristic  songs.  I  found  this  am- 
phibian very  common  in  small  ponds  at  the  roadside,  and  there  were  scores  of 
egg  masses,  attached  in  most  cases  to  blades  of  grass.  No  tadpoles  had  yet  ap- 
peared. So  numerous  were  the  egg  masses  that  I  collected  a  representative  lot 
of  them.  The  number  of  eggs  in  the  twenty-two  egg  masses  taken  were  as  fol- 
lows :  66, 45, 53, 33, 65, 46, 88, 38, 40, 67, 32, 50, 64, 87, 77, 15, 65, 51, 73, 45, 130,  and 
190.  The  number  of  eggs  here  found  is  much  greater  than  that  typical  of  the 
species  as  given  by  Dickerson  in  The  Frog  Book,  i.e.,  5-20  (page  159).  The 
masses  containing  130  and  190  eggs  respectively  seem  extreme.  There  is  a  possi- 
bility that  two  egg  masses  became  fused  in  these  cases,  but  except  for  the  fact 
that  the  numbers  of  eggs  are  unusually  high  there  is  no  reason  for  believing  so, 
as  the  gelatinous  material  remained  in  one  compact  body,  and  gave  no  evidence 
of  a  multiple  origin"  (H. }.  Pack,  Utah,  1920,  p.  7). 

"Eggs  and  adults  of  this  little  frog  were  taken  in  temporary  pools  formed  by 
the  melting  snow  along  a  railroad  right-of-way  near  Boulder,  during  the  first 
ten  days  of  May,  1914.  On  May  9  eggs  were  collected  in  the  four<ellcd  stage 
and  kept  out  of  doors  in  water  from  the  pool  in  which  they  were  found.  The 
development  of  these  eggs  was  very  rapid,  a  fact  which  may  be  correlated 
with  the  use  of  temporary  pools  as  the  spawning  grounds  by  this  species.  On 
the  nth  all  the  eggs  were  in  the  elongated  stage  preceding  hatching,  and  dur- 
ing the  1 2th  most  of  the  eggs  hatched.  The  tadpoles  of  Chorophiltts  trisenatus 
immediately  after  leaving  the  eggs  were  very  black  and  about  8  mm.  in  length, 
resembling  the  tadpoles  of  the  common  toad  in  outline"  (M.  M.  Ellis  and 
J.  Henderson,  Colo.,  1915,  p.  257). 

"May  17,  1919,  Fort  Sheridan,  111.  Found  Pseudacris  jenarum  \trisenata\ 
breeding  in  trenches  on  the  Rifle  Range  and  collected  a  couple  of  egg  clusters. 
The  eggs  are  attached  to  grass  stems  arranged  in  a  layer  one  or  two  eggs  thick 
all  around  the  stem,  forming  a  mass  about  two  inches  long  and  less  than  half 
an  inch  in  diameter.  .  .  .  June  22.  Tadpoles  of  this  species  are  metamorphosing 
in  the  same  pools  on  the  Rifle  Range  where  eggs  were  found  on  May  17.  They 
were  in  all  stages  of  development,  from  tadpoles  two-thirds  grown,  with  short 
hind  legs,  to  young  frogs  with  tails  nearly  absorbed.  The  measurements  of  sev- 
eral specimens  follow :  Two  full-grown  tadpoles  with  short  hind  legs  measured 
26  and  26.5  mm.  Four  with  large  hind  legs  and  fore-legs  almost  ready  to  burst 
out;  25.5, 27,  27.5,  and  28  mm.  Four  young  frogs  with  all  four  legs  well  devel- 
oped and  tails  in  different  stages  of  absorption  measured  10, 10, 10,  and  n  mm. 
in  body  length"  (P.  H.  Pope,  111.,  1919,  pp.  83-84). 

"Its  eggs  were  found  in  a  small  pool  on  the  22d  of  March.  They  were  at- 
tached to  twigs  in  small  and  large  bunches.  Each  egg  was  one-third  inch  in 
diameter,  including  the  usual  coating  of  jelly.  .  .  .  The  tadpoles  were  set  free 
on  April  5.  They  are  slenderer  than  the  larvae  of  the  Leopard  Frog,  and  not  so 


TREE  FROGS:  HYLIDAE  257 

dark  in  color.  They  arc  dark  gray  rather  than  black.  .  .  .  The  rudiments  of 
the  hinder  limbs  appear  about  the  2oth  of  April.  At  the  same  time  two  rows  of 
horny  teeth  appear  on  each  lip,  and  a  few  days  later  an  additional  row  on  the 
lower  lip.  These  teeth  are  minutely  denticulated  at  their  tips,  and  they  form 
an  admirable  apparatus  for  scraping  off  the  layer  of  nutritious  slime  that  covers 
all  objects  in  the  water.  There  are  from  55-95  of  these  teeth  in  each  row.  .  .  . 
By  April  20  the  length  has  become  one-half  inch  and  by  May  4  about  three 
quarters.  The  body  is  of  a  dark  color,  adorned  with  numerous  blotches  of  gold. 
The  belly  is  nearly  covered  with  a  shimmer  of  gold  and  coppery.  When  three- 
fourths  inch  long  the  young  were  observed  to  come  to  the  surface  and  take  in 
air.  By  the  2oth  of  May  the  young  have  attained  the  total  length  of  a  little  over 
an  inch.  By  the  26th  of  May  the  tadpoles  had  attained  a  length,  in  some  cases, 
of  28  mm.,  16  of  which  is  tail"  (O.  P.  Hay,  Ind.,  1889,  p.  773). 

"Many  of  them  about  this  time  succeed  in  releasing  their  forelegs  from  tfie 
skin  which  has  held  them  down.  Now  the  tadpoles  grow  smaller  instead  of 
larger.  This  is  largely  due  to  the  shortening  of  the  intestine  at  this  period  of 
transformation.  These  four-legged  tadpoles  are  very  lively  and  very  timid. 
They  show  a  great  inclination  to  get  out  of  the  water  and  to  hop  about.  They 
soon  lose  their  skill  in  swimming,  and  if  confined  to  the  water  too  long  will 
drown.  The  disks  are  seen  on  their  fore  feet  as  soon  as  these  feet  appear.  The 
tails  are  rapidly  absorbed,  and  by  the  i2th  of  June  all  have  become  little  frogs 
like  the  adults,  except  in  size,  At  the  time  of  transformation  the  length  of  the 
head  and  body  is  less  than  one-half  inch"  (same,  p.  471). 

"In  Kansas  the  eggs  arc  more  numerous  in  their  respective  masses  than  these 
previous  accounts  would  indicate,  numbers  varying  from  110-300,  the  mode 
being  about  140.  Wright  and  Wright  [Gen.,  1924,  p.  381]  give  500-800  as  the 
egg  complement.  One  female  observed  by  the  writer  laid  1,459  eggs  during  a 
single  night  in  the  aquarium  in  which  she  was  isolated.  The  membranes  of 
the  eggs  from  females  near  Lawrence  differ  somewhat  from  the  type  figured 
by  Wright  and  Wright  [Gen.,  1924,  pi.  i,  fig.  7],  The  outer  envelope  is  typi- 
cally about  3.0  mm.;  there  was,  in  several  bunches  of  eggs,  a  second  membrane 
about  2.1  mm.  in  diameter;  the  vitellus  is  as  given  by  Wright  and  Wright  (0.9 
to  1.2  mm.).  .  .  .  There  is  always  a  vitelhne  membrane  closely  applied  to  the 
vitellus"  (H.  M.  Smith,  Kans.,  1934,  PP-  4^7>  468). 

In  1936  K.  A.  Youngstrom  and  H.  M.  Smith  (Kans.,  1936,  pp.  629-630)  de- 
scribed the  tadpoles  of  this  species.  The  distinctive  thing  they  showed  is  lower 
labial  tooth  rows  3  instead  of  2.  This  is  a  fact  that  like  many  others  we  dis- 
covered soon  after  our  1929  paper. 

Journal  notes:  Ithaca,  N.Y.  Frogs  from  Buffalo,  N.Y.  On  the  morning  of 
April  2, 1907,  the  25  males  were  placed  with  the  5  females.  At  9:50  A.M.  the  first 
pair  was  recorded  and  within  20  minutes  the  female  began  laying.  With  one 
exception,  she  chose  a  different  perch  for  each  egg-laying  period,  thus  giving  a 
bunch  to  each  period  of  sexual  activity.  In  one  instance  she  sought  the  same 


258  HANDBOOK  OF  FROGS  AND  TOADS 

perch  three  successive  times,  This  pair  consumed  2%  hours  in  laying  a  com- 
plement of  500-600  eggs.  The  process  required  about  90  fertilizations  and  emis- 
sions. Each  time  from  two  to  ten  eggs  were  voided,  being  emitted  in  small 
strings,  a  condition  readily  seen  when  occasionally  the  eggs  were  unattached 
and  hung  down  from  the  vent  of  the  female.  All  the  eggs  were  laid  in  water 
66°F. 

April  4,  1928,  Hilton  Beach,  Hilton,  N.Y.  The  common  name,  "Chorus 
frog,"  is  very  appropriate.  Only  a  few  toads,  meadow  frogs,  and  peepers  were 
speaking.  It  was  a  very  deafening  chorus,  not  like  Hyla  femoralis  and  some 
larger  Hylas  but  very  pronounced.  This  Swamp  Cricket  Frog  may  be  in 
swampy  places  where  there  is  brush,  but  it  can  hardly  be  kept  down  in  note  to 
a  cricket  when  in  chorus. 

In  New  York  I  had  never  heard  it  except  at  Hilton,  Hilton  Beach,  Buffalo, 
and  Hamburg.  By  its  chorus,  we  found  it  in  Greece,  April  4,  at  Hilton  and 
Hilton  Beach  April  4  and  5,  and  at  Buttonwood  Creek  at  Parma  and  Greece 
town  line.  On  April  5  heard  it  near  Rochester,  near  Webster,  Williamson,  near 
Westbury,  3  miles  north  of  Auburn  on  the  Auburn  Weedsport  road. 

April,  1928,  Hilton,  N.Y.  West  of  Mt.  Reed  Church,  Greece,  we  found  it  m 
a  pond  in  an  orchard  where  several  small  trees  and  bushes  were,  and  also  brush. 
The  frogs  seemed  to  be  in  a  grassy  field,  but  mostly  along  a  fence-row  pond 
with  trees  and  brush.  Along  Parma  and  Greece  town  line,  they  were  in  over- 
flow pools  of  Buttonwood  Creek.  Not  brushy  there.  They  were  also  in  pools 
in  grassy  fields  toward  Braddock's  Bay. 

May  30,  1942.  With  Wilmcr  Webster  Tanner  went  i  mile  west  of  Provo, 
Utah,  to  a  spring.  Here  heard  a  great  chorus  of  P.  n.  triseriata.  Anna  caught  a 
female. 

June  8,  Flagstaff,  Ariz.  Went  to  Museum  of  Northern  Arizona.  Met  E.  D. 
McKee,  H.  S.  Colton,  Major  Brady.  At  museum  saw  Pseudacns  n.  tnsenattis 
from  Apache  County.  .  .  .  Tonight  went  to  Mary's  Lake,  7  miles  long.  At 
upper  end  a  great  chorus  of  P.  n.  tnsenatus.  When  we  returned,  in  a  stream 
back  of  the  motel  heard  several  P.  n.  triseriata. 

Authorities9  corner: 

F.  W.  Cragin,  Kans.,  1876-81,  p.  121.        A.  R.  Cahn,  111.,  1926,  pp.  107-108. 
B.  W.  Evermann  and  H.  W.  Clark,      F.  G.  Evenden,  Jr.,  Ida.,  1946,  p.  257. 

Ind.,  1920,  pp.  635-636. 

Florida  Chorus  Frog,  Florida  Winter  Frog 

Pseudacns  nigrita  verrucosa  (Cope).  Plate  LIV;  Map  17. 

Range:  "Peninsular  Florida.  Known  from  the  following  counties:  Alachua, 
Duval,  Volusia,  Marion,  Putnam,  Sumter,  Lake,  Hernando,  Pinellas,  Polk, 
St.  Lucie,  Charlotte,  Lee,  Collier,  Broward  and  Dade"  (Carr,  Fla.,  1940,  p.  56). 

Habitat:  Flatwoods,  prairie  lands,  glade  depressions,  and  ponds.  "Swamps 


TREE  FROGS:  HYLIDAE 


259 


Plate  LIU.  Pseudacris  nigrita  triseriata  Plate  LIV.  Pseudacris  nigrita  verrucow 

(XO-  i*3i5/>-  Females.  2.  Male.  4.  Eggs.  (X0«  l~^>  Males. 


260  HANDBOOK  OF  FROGS  AND  TOADS 

and  grassy  ponds  and  ditches;  moderately  common"  (O.  C.  Van  Hyning,  Fla., 


Size:  Adults  %-il/z  inches  (22-30  mm.  or  more). 

General  appearance  and  structure:  'The  length  of  the  head  to  the  posterior 
margin  of  the  membranum  tympani  enters  the  total  length  to  the  vent  three 
and  one-sixth  times.  The  head  itself  is  narrow  and  acuminate,  the  muzzle  pro- 
jecting acutely  beyond  the  labial  margin.  The  external  nares  mark  two-fifths 
the  distance  from  the  end  of  the  muzzle  to  the  orbital  border.  The  membranum 
tympani  is  only  one-fourth  the  diameter  of  the  orbit.  The  canthus  rostrahs  is 
distinct,  but  obtusely  rounded.  The  vomerine  fasciculi  are  approximated,  and 
near  the  line  of  the  posterior  border  of  the  nares,  which  are  larger  than  the 
minute  ostiu  pharyngca.  The  tongue  is  large  and  wide  behind  and  faintly 
emarginate. 

"The  heel  of  the  extended  hind  limb  extends  to  between  the  orbit  and  nos- 
tril: the  femur  is  short,  while  the  tarsus  is  long,  a  little  exceeding  half  the 
length  of  the  tibia,  and  exceeding  the  length  of  the  remainder  of  the  foot, 
minus  the  longest  toe.  The  skin  of  the  gular  and  sternal  region  is  smooth;  of 
the  abdomen,  areolate.  That  of  the  dorsal  region  is  tubercular,  smooth  warts  of 
large  and  small  size  being  irregularly  crowded  over  its  entire  surface,  and  not 
at  all  resembling  the  areolate  surface  of  the  belly, 

"Color  above  leaden,  with  three  longitudinal  rows  of  darker,  light-edged 
spots,  extending,  one  on  each  side,  and  one  on  the  median  line.  They  are  each 
composed  of  a  series  of  spots  joined  end  to  end.  Femur  and  tibia  cross-barred. 
Upper  lip  dark  plumbeous,  with  a  series  of  five  white  spots;  a  similar  spot  be- 
low the  tympanum.  Inferior  surfaces  yellowish.  .  .  . 

"From  Volusia,  Florida;  Mrs.  A.  D.  Lungren.  'This  Chorophtlns  is  similar 
in  proportions  to  the  C.  triseriatus,  but  is  well  distinguished  by  the  characters 
of  the  skin,  and  the  coloration.  The  tubercular  upper  surface  is  quite  peculiar, 
and  the  smooth  gular  region  is  equally  wanting  to  the  Northern  Frog.  The 
dorsal  skin  is  somewhat  like  that  of  Acns  gryllus  "  (E.  D.  Cope,  Fla.,  1877, 

PP.  87-88). 

"Cope  is  clearly  in  error  in  stating  |Gcn.,  1889,  p.  338]  that  verrucosus 
differs  from  C.  nigritus  (Le  Conte)  in  the  somewhat  longer  tarsus,  and 
from  C.  jenarum  (Baird)  in  the  longer  hind  leg.  He  records  the  tarsus  of 
nigritus  as  9.4  mm.,  and  the  'hind  leg'  of  jeriamm  as  1.77  in.  (45  mm.),  as  com- 
pared with  a  'hind  limb1  of  26  mm.  in  verrucosus.  The  type  of  verrucosus  is 
unquestionably  immature"  (M.  K.  Brady  and  F.  Harper,  Fla.,  1935,  p.  108). 

These  authors  emphasize  "the  maxillary  stripe  broken  to  form  a  series  of 
irregular  dots  and  bars  or  lacking"  (A.  F.  Carr,  Jr.,  Fla.,  1934,  p.  32). 

Voice:  "During  the  night  of  January  20,  1932,  R.  F.  Deckert  and  I  were  col- 
lecting in  the  Royal  Palm  Hammock,  on  Paradise  Key.  In  this  tropical  setting 
we  were  surprised  to  hear  the  trilling  of  Pseudacris  coming  from  the  'glade'  on 


TREE  FROGS:  HYIJDAE  261 

the  west  side  of  the  hammock.  The  call,  though  possibly  a  little  higher  in  pitch, 
was  remarkably  similar  to  that  of  P.  n.  feriarum  as  heard  in  the  vicinity  of 
Washington,  D.C.  The  chorus  was  very  small  and  localized.  After  considerable 
difficulty  we  managed  to  approach  the  position  of  the  calls  and  found  them 
coming  from  depressions  and  'potholes'  among  the  limestone  rocks  and  'glade* 
vegetation.  These  depressions  contained  shallow  ponds  that  were  choked  with 
hnardia.  Once  we  had  gotten  into  their  immediate  vicinity,  the  frogs  became 
silent,  but  by  running  a  finger  along  the  teeth  of  a  pocket  comb,  we  were  able 
to  imitate  the  voice  sufficiently  to  induce  them  to  call  again.  We  thus  located 
the  males,  which  occupied  well-separated  stations  and  were  sitting  on  projec- 
tions of  the  limestone  above  the  water  line  of  the  little  ponds"  (M.  K.  Brady 
and  F.  Harper,  Fla.,  1935,  p.  109). 

"The  call  is  similar  to  that  of  nigrita,  but  the  individual  crepitations  come 
much  faster — perhaps  twice  as  fast — and  to  some  extent  suggest  the  song  of 
fmarum"  (A.  F.  Carr,  Jr.,  Fla.,  1940,  p.  56). 

Breeding  and  Journal  notes:  "Flat woods;  prairie  lands  of  the  south-central 
peninsula.  Not  common.  Habits  apparently  similar  to  those  of  nigrita.  Breed- 
ing February  to  August  15,  in  cypress  ponds,  flooded  meadows,  drainage 
ditches  in  the  muck  land  around  Lake  Okecchobce,  and  in  'pot-holes*  and 
ditches  in  the  South  Florida  limestone"  (A.  F.  Carr,  Jr.,  Fla.,  1940,  p.  56). 

On  Feb.  7,  1932,  O.  C.  Van  Hyning  of  Gainesville,  Fla.,  wrote  as  follows: 
"I  went  out  into  the  middle  of  the  Prairie  a  few  nights  ago,  where  there  was 
a  little  water,  and  caught  eight  males  and  one  female  of  l\  nigrita;  the  female 
and  one  male  I  watched  from  the  time  they  were  about  a  foot  apart,  until  they 
mated,  and  would  have  stayed  on  another  hour  or  so,  but  the  mosquitoes  had 
just  about  eaten  me  up,  so  I  caught  them.  They  have  so  far  failed  to  mate  again 
although  two  males  were  together  once.  The  amplexation  was  axillary.  I  am 
going  out  to  the  same  place  as  soon  as  I  finish  this  letter,  and  see  if  I  can  get  any 
additional  females  and  see  the  eggs  deposited." 

The  following  week,  on  Feb.  15,  he  wrote:  "I  have  .  .  .  two  pairs  mated, 
one  of  a  male  and  a  female,  one  of  a  female  and  two  males,  and  one  case  of  two 
males  [in  the  laboratory].  I  have  one  bunch  of  eggs  laid  in  the  laboratory,  but 
they  were  pretty  well  walked  over  before  I  rescued  them.  I  didn't  know 
whether  they  were  fertile  or  not  and  have  preserved  them." 

Brady  at  Royal  Palm  Hammock  wrote:  "In  addition  to  a  series  of  six  calling 
males,  we  took  a  pair  in  amplexus.  The  female  deposited  160  eggs,  a  few  sep- 
arately, but  the  majority  in  a  loose  mass  not  differing  from  the  egg  mass  of 
feriarum.  The  brown  and  white  eggs  resembled  those  of  jeriamm,  but  seemed 
somewhat  smaller,  the  vitellus  measuring  less  than  0.5  mm.  The  single  enve- 
lope measured  1.5-2  mm.  Development  was  very  rapid  and  hatching  took  place 
within  60  hours"  (M.  K.  Brady  and  F.  Harper,  Fla.,  1935,  p.  109). 

On  Feb.  4, 1933,  at  Gainesville,  Fla.,  Messrs.  A.  F.  Carr  and  H.  K.  Wallace 


262  HANDBOOK  OF  FROGS  AND  TOADS 

found  this  species  breeding  at  the  same  time  as  P.  ocularis  and  P.  ornata.  The 
eggs  they  sent  us  were  brown  and  cream  or  white,  the  vitellus  ^5  inch  (0.9-1.0 
mm.),  the  single  envelope  Ko-%  inch  (2.6-2.8  mm.). 

Cope  emphasized  the  warty  upper  surface.  We  have  seen  very  warty  ones 
in  the  southern  half  of  the  peninsula,  and  W.  J.  Hamilton,  Jr.,  at  Ft.  Myers 
March  n,  1940,  took  very  tubercular  specimens. 

Authorities9  corner: 
A.  F.  Carr,  Jr.,  Fla.,  1940,  p.  55. 

Western  Chorus  Frog 
Pseudacris  occidentalis  (Baird  and  Girard).  Plate  LV. 

This  frog  probably  does  not  exist. 

If  Cope  (1889)  was  unwarranted  in  assigning  Chorophilus  occidentalis 
(B  &  G)  (1853)  to  the  eastern  Rana  ornata  Holbrook  (1836)  or  Cystignathus 
ornatus  Holbrook  (1842),  what  then  is  this  Lttoria  occidentals  which  Baird 
and  Girard  described  in  a  paper  entitled  "A  List  of  Reptiles  Collected  in  Cali- 
fornia by  Dr.  John  L.  Le  Conte  .  .  ."?  We  prefer  to  believe  it  was  collected 
in  California.  All  the  other  batrachians  of  the  report  were  strictly  western 
forms.  Baird  and  Girard  report  Hyla  regilla  in  this  same  paper  (Calif.,  1853, 
p.  301),  first  describe  it  in  another  place  of  this  same  volume  (p.  174),  and  Hal- 
lowell  also  describes  Hyla  regtlla  in  the  same  volume  (p.  183)  as  Hyla  scapu- 
laris.  Nevertheless  the  Litona  occidentalis  of  this  report  might  well  be  another 
variant  specimen  of  the  very  variable  Hyla  regilla.  At  that  period  we  have  Hal- 
lowell  in  the  one  paper  (Calif.,  1856,  pp.  96-97)  describing  Hyla  regilla  three 
times  as  Hyla  scapulans,  Hyla  nebulosa,  and  Hyla  scapulans  var.  hypochondri* 
aca.  We  therefore  suspect  Litona  occidentalis  of  being  a  synonym  of  a  western 
Hyla,  presumably  Hyla  regilla.  A  glance  at  the  plate  of  Hyla  regilla  (Plate 
LXX)  will  show  at  once  how  many  resemblances  this  species  has  to  Pseudacris. 

In  1889  Cope  (Gen.,  p.  337)  speaks  of  C.  occidentals  (which  we  now  term 
P.  ornata)  as  ranging  "to  the  Wichita  River,  in  north  central  Texas.  Specimens 
were  sent  me  from  the  latter  locality  by  that  excellent  naturalist,  Jacob  Boll,  of 
Dallas.  ...  It  does  not  occur  in  California  as  supposed  when  first  described." 
What  are  these  Texan  forms? 

The  records  of  three  friends,  C.  S.  Brimlcy,  J.  Hurter,  and  J.  K.  Streckcr,  arc 
yet  to  be  discussed.  Brimley  (Ala.,  1910,  p.  n)  secured "Chorophilus  occidenta- 
Us  from  Bay  St.  Louis,  Mississippi,  February  and  April  1898,  5"  and  Choro- 
philus ornatus  "Hastings,  Florida,  June,  1901,  i;  Green  Cove  Springs,  Florida, 
July,  1898,  5."  Doubtless  a  specimen  in  U.S.  National  Museum,  no.  29189,  la- 
beled C.  occidentalis  Hastings,  Fla.  H.  H.  &  C.  S.  Brimley,  is  the  one  specimen 
Brimley  rightly  called  C.  ornatus  above.  He  is  the  only  person  to  receive  each 
from  the  Southeast  and  the  only  one  in  that  period  to  call  the  Floridan  form 
C.  ornatus.  But  what  was  the  Chorophilus  occidentalis  of  Bay  St.  Louis,  Miss.? 


TREE  FROGS:  HYLIDAE  263 

In  the  Amphibians  and  Reptiles  of  Arkansas,  Hurter  and  Strecker  record 
Chorophilns  occidentalis  from  "Hot  Springs  (Combs)."  Strecker  knows 
Pseudacris  strecJ^eri  (ornata  of  Cope,  1889)  and  has  contributed  as  much  as  or 
more  than  anyone  living  to  its  habits,  but  the  Pseudacris  occidentalis  is  the 
puzzle. 

Since  the  paragraph  above  was  written,  there  have  come  to  hand  Strecker's 
records  of  Pseudacris  ornata  (strecl^eri)  and  Pseudacris  occidentalis.  In  recent 
times  only  two  people  have  collected  Pseudacris  occidentalis  in  life  to  recog- 
nize it.  Richard  F.  Deckert  secured  it  in  Jacksonville  along  with  P.  n.  mgnta 
and  P.  octtlaris.  This  author  does  not  record  P.  ornata  and  his  descriptions  of 
P.  occidentalis  are  those  of  P.  ornata,  but  the  records  of  the  other  author  who 
has  recently  seen  this  species  cannot  be  so  readily  allocated  to  P.  ornata  (occi- 
dentals of  Cope  and  Deckert),  or  P.  strec\en  (ornata  of  Cope  and  Strecker). 
The  only  collector  in  America  who  has  taken  P.  occidentalis  and  the  Texan 
P.  ornata  (now  called  P.  strec\eri)  in  the  same  region,  alive,  is  J.  K.  Strecker. 

On  Sept.  i,  1929,  he  wrote  of  the  capture  of  specimens  of  each  species  in  the 
Fort  Worth  region: 

"Western  Chorus  Frog  (Pseudacris  occidentalis  (Baird  and  Girard)).  I  cap- 
tured two  specimens.  One,  a  junior,  was  caught  four  miles  beyond  Benbrook, 
in  grass  under  the  edge  of  a  rock  along  a  roadside;  the  second  example,  an 
adult,  was  caught  among  grass  growing  along  a  creek  bank.  Ornate  Chorus 
Frog  (Pseudacris  ornata  Holbrook).  North  and  east  of  Fort  Worth,  I  caught 
three  adults  among  grass  growing  on  the  banks  of  small  creeks"  ( J.  K.  Strecker, 
Jr.,  Tex.,  1929,  p.  n). 

In  one  museum  some  little  P.  octtlaris  were  termed  P.  occidentals.  Of  several 
different  museum  accessions  considered  P.  occidentalis,  we  have  made  the 
positive  note:  "This  is  P.  strecJ^eri.  Possibly  some  of  the  Texan  representatives 
may  prove  P.  n.  clarfyi" 

In  1930  we  visited  Dr.  J.  K.  Strecker,  Jr.,  and  received  many  kindnesses  from 
him.  While  we  were  in  the  field  for  P.  strec\eri  (ornata  of  Strecker),  he  re- 
marked that  what  he  had  recorded  as  P.  occidentalis  appeared  different  from 
jP.  strecljfri  (ornata)  and  had  different  habits.  He  kindly  loaned  us  what  speci- 
mens he  had  of  P.  occidentalis.  At  first  they  puzzled  us,  but  the  more  we  ex- 
amined them  the  more  we  inclined  to  interpret  them  as  the  spotted  individuals 
of  the  so-called  P.  n.  clarfyi.  Taking  his  Nos.  4501  and  4502  (Baylor  Museum) 
of  Fort  Worth  and  Bluff  Creek,  McLennan  Co.,  and  comparing  them  with  his 
3507  P.  nigrita,  we  call  all  three  P.  n.  clar\ii.  Another  specimen,  P.  occidentals, 
from  Gayle,  La.  (Frierson),  may  be  a  large  female  P.  nigrita  (triseriata) . 

Before  we  saw  these  specimens  (4501,  4502)  we  wondered  if  they  might  be 
like  P.  strec\eri,  but  in  longer  head-to-tympanum,  wider  head,  smaller  eye, 
wider  internasal,  shorter  hind  limb,  shorter  first,  second,  third,  fourth,  and  fifth 
toes,  P.  strec^eri  does  not  overlap  the  relative  measurements  (number  of  times 
in  length)  of  these  two  P.  occidentalis  specimens. 


264  HANDBOOK  OF  FROGS  AND  TOADS 

45OI>  4502 

Head  to  tympanum  3.16-  3.29  2.38  -  3.0 

Width  of  head  3.11-3.47  2.4    -2.88 

Internasal  sp.  11.2-11.8  8.0    -10.2 

Eye  8.44-  9.33  9.33  -12.0 

Hind  limb  0.66-  0.72  o-734~  0-87 

First  toe  5.45-  5.6  6.22  -  9.0 

Second  toe  4.3  -  4.52  4.66  -  5.53 

Third  toe  2.8  -  2.95  3.11  -  3.6 

Fourth  toe  2.0  -  2.18  2.15  -  2.57 

Fifth  toe  2.66-  2.68  2.8    -  3.5 

The  relative  measurements  of  Nos.  4501  and  4502  fall  within  the  relative 
measurements  of  P.  n.  dartyi  in  almost  every  respect  except  head  length  and 
intertympanic  and  internasal  widths.  Most  of  the  measurements  of  No.  1378, 
the  specimen  from  Gayle,  La.,  fall  within  those  of  P.  n.  triscnata. 

All  in  all  our  conclusions  are  that  these  P.  occidentalis  specimens  may  be 
forms  of  P.  nigrita  and  are  not  P.  streckfn  (ornata  of  Strecker  and  Cope). 
Frogs  called  P.  occidentalis  in  Georgia  and  Florida  are  P.  ornata  (Holbrook). 

Little  Chorus  Frog,  Swamp  Tree  Frog,  Swamp  Cricket  Frog,  Tree  Frog, 
Savanna  Cricket  (Bartram),  Least  Swamp  Frog 

Psetidacns  ocularis  (Holbrook).  Plate  LVI;  Map  18. 

Range:  North  Carolina  to  southern  Florida. 

Habitat:  Sphagnum  edge  of  cypress  pond.  Grassy  or  sedgy  area  of  pond  or 
wet  edge  of  pine  barrens. 

May  i83  1921,  Pseudacris  ocularis  may  start  up  from  sphagnum  edge  of 
cypress  pond  or  may  start  up  from  the  ground.  They  will  leap  1-1%  feet  at 
times.  These  specimens  sat  on  Eriocaitlon,  small  Sarracema  minor,  sedge,  and 
saw  palmetto.  We  took  a  dozen  of  them.  On  June  25,  we  found  them  common 
in  one  pond,  many  in  bamboo  (smilax)  vines,  on  bushes  some  4-5  feet  up, 
others  on  level  of  water.  July  15,  we  found  them  abundant  in  outer  edges  of 
cypress  ponds,  margins  of  cypress  bays.  Most  of  them  we  caught  in  the  grass 
near  the  edge  of  ponds.  June  17, 1922,  we  found  them  in  grassy  and  sedgy  areas 
on  wet  edge  of  pine  barrens  or  outer  edge  of  cypress  bay  or  branch. 

"Wet  grassy  places,  especially  in  the  pine  flatwoods,  common"  (O.  C.  Van 
Hynmg,Fla.,  1933,  p.  4). 

Size:  Adults,  %-%  inch.  Males,  11.5-15.5  mm.  Females,  12.0-17.5  mm. 

General  appearance:  These  are  the  brownies  of  frogdom  in  the  United 
States.  They  may  be  uniform  gray,  brown,  greenish,  or  reddish  on  the  back 
with  a  dark  vitta  from  the  eye  backward  as  a  stripe  of  variable  length.  This  is 
set  off  by  a  light  area  below  the  eye  extending  backward  to  the  shoulder.  There 
may  be  a  dark  triangle  between  the  eyes  with  a  stripe  extending  down  the 


TREE  FROGS:  HYLIDAE 


265 


iP;-:';fr  "J-  ^:'-'"C;-^^ 
sis^^d^i^1.^1 ! 

|iP^'>X--:-:- 


Plate  LV.  Pseudacris  occidcntalis  (X%)- 
1-7.  Males.  8.  Female. 


Plate  LVI.  Pseudacris  ocularis.  1,4,5,7,8. 
Adults  (XiVO-  a/>-  Males  (Xi%)-  3-  Male 
croaking  (XO- 


266  HANDBOOK  OF  FROGS  AND  TOADS 

mid-back  and  a  stripe  on  either  side  of  this.  These  little  frogs  are  so  tiny,  so 
delicate,  that  it  does  not  seem  possible  that  they  are  adult  frogs.  Their  form  is 
slender,  their  legs  long,  their  eyes  bright  and  beadlike,  their  snouts  very  pointed 
and  extending  beyond  the  lower  jaw.  The  nostrils  are  on  the  sides  of  the 
pointed  snout.  These  little  midgets  can  turn  their  heads  or  tip  them  upward 
or  sideways  without  turning  the  body. 

Color:  On  May  17, 1921,  Okefinokee  Swamp,  Ga.,  we  had  diverse  colorings 
in  a  lot  of  16  in  one  pan.  Some  were  tawny,  hazel,  or  amber  brown  on  upper 
surfaces;  others  buckthorn  brown;  one  a  light  grayish  olive  or  light  olive-gray; 
another  deep  olive-gray.  The  stripe  on  the  upper  jaw  was  almost  cream,  chal- 
cedony yellow,  or  glass  green.  Belly  chamois  or  cream-buff.  Iris  cinnamon. 

On  Aug.  1 8, 1922,  at  Milliard,  Fla.,  found  a  little  Psettdacris  ocularts  which 
is  cinnamon  or  ochraceous-tawny  in  color.  It  is  an  adult  male  with  a  plaited 
throat.  The  ten  specimens  vary  in  color  from  gray,  greenish,  light  yellowish 
olive  or  dull  citrine  through  to  chestnut  or  bay;  four  specimens  were  of  the 
yellowish  olive  or  gray  group,  four  of  the  chestnut  or  bay  group,  and  two  in- 
termediate. The  most  distinctive,  yet  not  always  constant,  mark  is  a  line  of 
dark  color  which  runs  from  the  snout  or  nostril  through  the  eye  and  tympanum 
to  or  beyond  the  arm  insertion.  In  three  specimens  it  runs  halfway  to  the  hind 
limbs.  In  two  more  it  just  passes  the  arm  insertion,  as  Holbrook  figures  it.  In 
two  this  dark  stripe  is  absent.  In  the  striped  individuals  the  limbs  bear  cross- 
bars. In  all  material,  the  entire  upper  surfaces  have  a  very  fine  speckling  of 
close-set  dark  dots.  The  under  parts  arc  whitish  or  yellowish  white  with  the 
fine  speckled  dots  of  the  upper  parts  all  over  the  ventral  side.  In  some  the  un- 
derside of  limbs  is  almost  as  dark  as  the  upper  parts. 

When  four  specimens  arrived  May  7, 1929,  from  Gainesville,  Fla.,  from  O.  C. 
Van  Hyning,  in  sphagnum,  all  were  different:  one  was  uniform  carnelian  red 
above  with  no  stripes  except  vitta  on  side;  another  dark  olive;  one  deep  olive- 
buff  with  three  stripes  on  back;  and  one  avellaneous.  Later  the  carnelian  red 
was  entirely  changed. 

Structure:  Midgets  in  size;  snout  pointed  and  projecting  beyond  the  lower 
jaw;  nostrils  on  sides  of  snout;  slender  in  form,  hind  legs  very  long;  eyes  bright 
and  beadlike;  disks  on  fingers  and  toes  small  but  distinct;  skin  of  back  covered 
with  very  fine  warts. 

"H.  ocularis.  H.  ocularis  Daudin,  i  c.  p.  32.  Hyloclcs  ocularts  Holbrook,  pi. 
35.  Above  brown  or  bronzed  or  silvery  grey,  very  finely  speckled  with  dusky 
or  darker,  a  tolerably  wide  band  of  black  proceeds  from  the  tip  of  the  nose  to 
the  middle  or  beyond  the  middle  of  the  sides,  this  is  bordered  beneath  with 
white.  Chin  and  underside  of  the  thighs  speckled  with  black.  Legs  speckled 
like  the  back  and  more  or  less  spotted  and  barred  with  dusky,  fingers  and  toes 
all  furnished  with  small  disks.  Length  .6.  Inhabits  Georgia.  The  smallest  of  all 
known  Ranina,  From  the  small  size  of  this  and  the  preceding  species,  the  web 


TREE  FROGS:  HYLIDAE 


267 


between  the  third  and  fourth  toes  is  not  very  perceptible"  (J.  Le  Conte,  Gen., 
1855,  p.  429). 

Voice:  The  call  is  a  high,  shrill,  cricketlike  chirp  or  trill.  Vocal  sac  of  the 
male  even  when  collapsed  sometimes  appears  to  cover  almost  half  of  the  venter 


/VX  Hyh  regilla 

.  wright or um 

Pseud acris  ocularis 

P.  n.  clarkii 
I".  P.  n.  sepfenfr/ona//s 

P.  n.  triseriata 


Map  18 

of  the  body.  The  rear  of  it  may  reach  caudad  to  the  pectoral  region,  to  the  line 
connecting  one  arm  insertion  with  the  other.  Our  first  acquaintance  with  a 
chorus  came  the  evening  of  May  20,  1921,  in  Okefinokee  Swamp,  Ga.  Went 
down  to  the  pond  east  of  the  Negro  quarters  and  it  sounded  as  if  bedlam  had 
broken  loose.  I  heard  a  cricketlike  note  everywhere.  It  was  Pseudacris  ocularis. 
One  frog  was  on  a  grassy  mat,  one  on  a  log,  another  calling  from  pine  brush 
at  the  edge  of  the  water,  another  on  the  bole  of  a  tree.  They  do  not  like  the 
electric  spotlight.  One  on  pine  brush  worked  up  into  the  leaves.  We  flashed  one 


2&I  HANDBOOK  OF  FROGS  AND  TOADS 

on  the  side  of  a  black  gum.  Its  throat  pouch  was  transparent  and  we  could  see 
through  it  and  discern  the  bark  behind.  The  croaks  were  given  30-65  per  min- 
ute. The  note  is  high-pitched,  penetrating,  can  be  heard  150-200  feet  away. 
Whether  characterized  as  chirp,  trill,  cheep,  squeak,  or  high  shrill  insect  call, 
it  surely  is  a  loud  piercing  call  for  so  little  a  mite  of  frog  flesh.  It  is  an  amusing 
little  creature  as  it  squeezes  its  slender  body  and  throws  out  its  large  sac  one- 
half  the  size  of  the  body. 

Breeding:  It  breeds  from  January  to  September.  The  brown  and  cream  eggs 
are  single,  laid  on  the  bottom  of  ponds  and  in  vegetation  in  shallow  water, 
about  100  in  number.  The  egg  is  Mo-Ho  inch  (0.6-0.8  mm.),  the  single  envel- 
ope ^o-Via  inch  (1.2-2.0  mm.).  The  greenish  tadpole  is  small,  !%«  inch  (23 
mm."),  its  tail  long.  Its  tooth  ridges  are  %.  After  45  to  70  days  the  tadpoles  trans- 
form, June  30  to  August  18,  at  '}io-%  inch  (7-9  mm.). 

Journal  notes:  In  1921,  on  July  3,  we  took  a  handcar  trip  to  Honey  and  Black 
Jack  Islands.  It  was  a  misty  day  at  first,  then  sunny  a  while;  it  rained  all  the 
afternoon,  however.  The  temperature  was  from  71° -76°.  On  the  sphagna- 
ceous  strand  of  Honey  Island  heard  these  creatures  near  Black  Jack  Island;  in 
mid-afternoon  on  Honey  Island,  in  Cypress  Bay  between  the  Pocket  and  Jones 
Island,  and  all  over  the  prairie,  in  sphagnum  of  wet  thickets  especially.  Also  all 
along  the  trestle  between  Jones  Island  and  Billy's  Island.  Never  heard  such  a 
frog  din. 

On  July  29, 1921,  we  were  caught  in  a  downpour  on  Black  Jack  Island.  We 
were  encamped  in  the  moist  pine  barrens.  All  about  was  the  strongest  chorus 
of  Psettdacris  ocularis  we  ever  heard.  Temperature  about  75°. 

In  1922,  we  secured  them  on  Chesser  Island,  near  old  Suwannee  canal,  along 
the  road  to  Folkston,  in  ponds  near  Trader's  Hill,  near  an  overgrown  sphag- 
num bog  outside  the  swamp,  and  along  the  road  from  Folkston,  Ga.,  to  Jack- 
sonville, Fla. 

Authorities'  corner: 

W.  Bartram,  Gen.,  1791,  p.  278.  J.  E.  Holbrook,  Gen.,  1842,  p.  138. 

F.  M.  Daudin,  Gen.,  1801-1803,  VIII,      G.  K.  Noble,  Gen.,  1923,  pp.  1-5. 

68-69.  F.  Harper,  Ga.,  1939,  p.  148. 

Ornate  Chorus  Frog,  Ornate  Tree  Frog,  Ornate  Swamp  Frog,  Swamp  Cricket 
Frog,  Ornate  Winter  Frog,  Holbrook  Chorus  Frog 

Pseudacris  ornata  (Holbrook).  Plate  LVII;  Map  16. 

Range:  North  Carolina  (B.  B.  Brandt,  1933)  to  Georgia,  Florida,  and  pos- 
sibly west  along  the  Gulf  to  Louisiana.  Alabama  (Viosca). 

Habitat:  Holbrook  wrote  of  its  habits:  "I  have  always  found  it  on  land  and 
in  dry  places,  and  frequently  in  cornfields  after  light  summer  showers.  It  is 
very  lively  and  active,  making  immense  leaps  when  pursued,  and  consequently 
is  taken  with  great  difficulty."  "The  structure  of  this  species  indicates  terres- 


TREE  FROGS:  HYLIDAE  269 

trial,  possibly  subterraneous  habits.  I  have  dug  specimens  out  of  the  sweet  po- 
tato hills  in  my  garden"  (R.  F.  Deckert,  Fla.,  1915).  M.  J.  Allen  found  it 
breeding  in  grass-land  pools.  "Grassy  ditches  and  ponds;  moderately  common" 
(O.  C.  Van  Hyning,  Fla.,  1933,  p.  4). 

Size:  Adults,  1-1%  inches.  Males,  25-35  mm-  Females,  28-36  mm. 

General  appearance:  This  is  a  large  Pseudacris,  looking  almost  like  a  small 
Rana  sylvatica.  The  general  color  of  this  frog  is  chestnut  brown  with  two  in- 
distinct darker  dorsal  bars  and  an  indistinct  darker  spot  between  the  eyes. 
There  is  a  prominent  dark,  almost  black,  vitta  from  tip  of  snout  through  eye 
over  tympanum,  usually  beyond  the  arm  insertion,  often  extending  halfway  to 
the  groin.  Two  frogs  with  more  prominent  stripes  have,  between  the  eyes, 
Y-shaped  triangles,  which  in  the  other  four  do  not  show.  The  groin  spots  are 
inclined  to  be  almost  black  oblique  bars,  with  light  outlines.  In  fact,  the  spots 
on  groin  are  on  a  yellow  background.  The  tiniest  rim  of  the  upper  jaw  is  dark, 
but  above  this  is  a  conspicuous  light  line  extending  to  the  arm  insertion  and 
broadest  under  the  eye.  On  the  front  of  the  upper  arm  is  a  dark  bar.  The  eyes 
are  very  prominent  with  the  upper  part  of  the  iris  a  bright  gold  band.  The 
throat  of  the  male  is  dark  olive,  with  a  central  longitudinal  plait.  The  throat  of 
the  female  is  light.  The  legs  are  barred. 

Cope  characterized  it  well  enough  but  added  to  the  confusion  by  calling  it 
Chorophtlus  occidentalis  B.  &  G. 

"Muzzle  rounded  in  profile,  projecting.  Skin  of  upper  surface  smooth.  .  .  . 
More  slender  [than  C.  ornatus\\  width  of  head  entering  length  3  to  3.5  times; 
nostril  nearer  end  of  muzzle  than  orbit;  posterior  foot  longer,  not  webbed,  and 
without  subarticular  tubercles;  heel  reaching  middle  of  orbit"  (E,  D.  Cope, 
Gen.,  1 889,  p.  332). 

G.  A.  Boulenger,  who  called  it  Chorophtlus  copti,  wrote:  "Vomerine  teeth 
behind  the  level  of  the  choanae;  tibio-tarsal  articulation  not  reaching  the  eye; 
back  immaculate"  (Gen.,  1882,  p.  332). 

ft  Chorophtlus  copii.  Tongue  circular,  entire  vomerine  teeth  in  two  slightly 
oblique  oval  groups  behind  the  level  of  the  choanae.  Head  slightly  longer  than 
broad;  snout  rounded,  as  long  as  the  diameter  of  the  orbit;  canthus  rostralis 
rather  indistinct;  interorbital  space  as  broad  as  the  upper  eyelid;  tympanum 
somewhat  more  than  half  the  diameter  of  the  eye.  Fingers  and  toes  moderate, 
the  tips  very  indistinctly  dilated;  first  finger  shorter  than  second;  toes  with  a 
slight  rudiment  of  web;  subarticular  tubercles  small;  no  outer  metatarsal  tu- 
bercle. The  hind  limb  being  carried  forwards  along  the  body,  the  tibio-tarsal 
articulation  marks  the  tympanum.  Skin  smooth  above,  granulate  on  the  belly. 
Olive  above;  sides  with  three  black  spots,  viz,  a  streak  from  the  eye  to  the 
shoulder,  an  oval  spot  in  the  middle  of  the  side,  and  one  or  two  smaller  ones 
on  the  loan.  Georgia.  A  9  Georgia. 

"Mr.  Cope  considers  this  frog  specifically  distinct  from  C.  ornatus  and  iden- 
tifies it  with  Hyla  ocularis  of  Daudin.  The  latter  opinion  I  cannot  share,  as 


270  HANDBOOK  OF  FROGS  AND  TOADS 

nothing  in  the  French  author's  description  and  figure  indicates  the  least  anal- 
ogy with  this  form.  The  toes  are  represented  as  half-webbed  in  H.  ocularis 
and  the  digital  expansions  of  C.  copii  are  so  slight  that  they  would  certainly 
have  escaped  Daudin's  observation.  Moreover  the  colour  is  quite  different" 
(same,  p.  334). 

Color:  Gainesville,  Fla.,  from  O.  C.  Van  Hyning,  March  24, 1932.  First  male. 
The  background  color  of  the  back  and  top  of  head  is  sayal  brown  or  tawny- 
olive,  becoming  on  top  of  fore  limb  and  eyelids  and  top  of  head  cinnamon  or 
orange-cinnamon.  The  hind  legs  may  be  verona  brown.  The  band  on  either 
side  of  middle  of  back  is  indistinct  and  is  snuff  brown  or  Saccardo's  umber. 
The  stripe  from  snout  obliquely  upward  to  nostril,  thence  through  eye  over 
tympanum,  shoulder  insertion,  two-thirds  of  way  to  groin,  may  be  black  or 
sepia  in  front  and  warm  sepia  at  the  caudal  end.  The  one  or  two  groin  spots 
may  be  warm  sepia,  chocolate,  or  walnut  brown.  The  three  bars  on  femur  are 
the  same  color.  The  interstices  of  groin  spots  are  light  cadmium  to  lemon  yel- 
low. There  are  also  flecks  of  the  same  on  the  front  of  the  upper  half  of  the  fe- 
mur. The  front  of  the  femur  is  partly  dark  vinaceous-brown.  The  rear  of  femur 
has  punctulations  of  vinaceous-brown  or  dark  vinaceous-brown.  Lower  end  of 
rear  of  femur  has  a  few  spots  of  lemon  yellow  or  lemon  chrome.  The  side  of 
body  is  vinaceous-buflf,  at  times  becoming  on  top  of  hind  legs  and  side  light 
drab  or  pale  drab-gray.  The  light  band  on  upper  edge  of  upper  jaw  is  pale  dull 
green-yellow,  becoming  at  angle  of  mouth  and  under  the  tympanum  white. 
The  dark  vitta  from  eye  backwards  on  its  upper  margin  has  a  thin  outline  of 
seafoam  yellow  to  white;  the  groin  spot  or  spots  have  a  suggestion  of  the  same 
thing.  The  upper  jaw  rim  is  a  thin  line  of  buffy  brown.  The  eye  color  in  the 
lower  half  partakes  of  color  of  the  vitta,  wherein  the  upper  half  is  the  color  of 
top  of  head.  There  is  a  prominent  natal  brown  or  vinaceous-brown  band  from 
the  lateral  vitta  down  across  the  arm  insertion  extending  even  onto  the  forearm 
to  make  its  first  crossbar.  On  forearm,  there  is  a  prominent  bar  of  same  color 
at  base  of  hand.  The  underside  of  fore  and  hind  limbs  is  brownish  vinaceous. 
The  belly  is  a  dirty  white  with  considerable  spotting  in  the  rear  pectoral  region 
of  drab  or  olive-brown.  The  throat  is  pronouncedly  discolored,  its  rear  sharply 
outlined  by  a  fold;  the  caudal  half  is  olive-ocher  or  old  gold,  the  cephalic  por- 
tion is  Saccardo's  olive,  the  two  central  plaits  becoming  olive-citrine.  The  edge 
of  the  lower  jaw  is  spotted  with  white  or  pale  dull  green-yellow.  After  we  had 
finished  describing  it,  the  dorsal  color  came  to  be  uniform  with  no  suggestion 
of  long  dorsal  stripes. 

Second  male.  A  uniform  mahogany  red,  chestnut,  or  bay  on  the  back. 

Female.  Charleston,  S.C.,  from  E.  B.  Chamberlain,  Nov.  4, 1932.  The  back- 
ground color  of  the  back  and  top  of  head  is  Hay's  russet,  becoming  on  side 
orange-cinnamon  or  sayal  brown.  The  two  parallel  dorsal  bands  are  liver 
brown  or  carob  brown,  so  also  are  the  bars  on  the  femur  and  tibia.  Those  on 
tarsus  and  foot  merge  into  black.  Snout  stripe,  eye  vitta,  lateral  and  groin  spots 


TREE  FROGS:  HYLIDAE  271 

are  light  cadmium  or  lemon  yellow.  The  rear  of  the  femur  much  as  in  the  male. 
The  light  band  along  the  upper  jaw  is  very  narrow  and  whitish.  The  upper  jaw 
rim  is  bone  brown.  The  eye  is  like  that  of  the  male.  The  venter  is  more  or  less 
like  that  of  the  male,  except  that  the  throat  is  white  without  discolor. 

Structure:  Form  more  slender  than  Pseud acris  strecf^eri;  head  narrower  than 
P.  stred(eri;  arms  and  legs  longer  and  more  slender  than  P.  strecl^eri;  fingers 
and  toes  long  and  slender,  with  mere  trace  of  web  at  base  of  toes;  snout  pointed ; 
nostril  equidistant  between  snout  and  eye;  vomerine  teeth  between  nares. 

"The  head  is  small,  with  a  broad,  indistinct,  triangular  spot  between  the  or- 
bits, the  apex  of  which  is  directed  backwards.  A  black  line  extends  from  the 
snout  to  the  orbit  of  the  eye,  including  the  nostrils;  below  this  black  line  is  a 
yellowish  blotch,  covering  most  of  the  upper  jaw.  The  lower  jaw  is  cinereous 
above  and  white  below.  The  mouth  is  small,  and  the  palate  is  armed  with  two 
groups  of  exceedingly  minute  teeth  between  the  posterior  nares.  The  nostrils 
are  placed  on  a  slight  prominence.  The  eyes  are  large  and  projecting,  the  pupil 
very  dark,  the  ins  of  a  golden  colour.  The  tympanum  is  small,  very  dark  col- 
ored, and  placed  in  a  dark  vitta,  or  blotch,  which  extends  behind  the  orbit  to 
within  a  short  distance  of  the  shoulder.  The  body  is  short,  of  a  delicate  dove- 
colour  above,  with  two  or  more  oblong  spots  of  dark  brown,  margined  with 
yellow,  on  each  side  of  the  vertebral  line;  below  these,  and  on  each  flank,  are 
three  smaller  spots,  likewise  margined  with  bright  yellow,  the  anterior  one 
being  the  largest;  these,  with  a  smaller  one  above  the  vent,  form  a  triangle  on 
each  flank.  .  .  .  The  toes  are  five  in  number,  not  palmated,  the  two  outer  ones 
only  are  united  at  the  base.  Dimensions.  Length  of  body  from  the  snout  to  the 
vent,  1%  inches;  of  the  thighs,  %  an  inch;  of  the  leg,  %  an  inch;  of  the  tar- 
sus and  toes,  nearly  %  of  an  inch"  (J.  E.  Holbrook,  Gen.,  1842,  pp.  103-104). 

Voice:  "This  call  is  very  loud,  similar  in  pitch  to  that  of  Hyla  picl(enngi,  but 
much  shorter,  and  at  a  distance  sounds  like  the  ring  of  a  steel  chisel,  when 
struck  by  a  hammer  .  .  ."  (R.  F.  Deckert,  Fla.,  1915). 

Norman  Davis  of  Gainesville,  Fla.,  says,  "The  call  reminds  me  of  Hyla  cru- 
cifer but  is  without  the  trill." 

"I  have  timed  ornata  and  found  the  calls  to  be  repeated  from  65-80  times  a 
minute.  I  have  never  actually  timed  crucifer,  but  its  calls  are  a  good  deal  slower, 
probably  40-50  per  minute.  Ornata 's  call  is  a  single  sharply  terminated  note, 
while  crucifer  s  is  a  more  deliberate  slur  from  a  lower  to  a  higher"  (letter  from 
A.F.Carr,Feb.5,i933). 

Breeding:  Winter  months.  November,  December,  January  to  March  depend- 
ing upon  the  rains.  H.  K.  Wallace  and  A.  F.  Carr,  Jr.,  at  Gainesville,  Fla.,  se- 
cured eggs  laid  Feb.  5, 1933.  They  are  brown  and  cream  or  white,  and  measure: 
the  vitellus  Vzs  inch  (0.9-1.0  mm.),  the  single  loose  envelope  /4-%  inch  (3.6- 
4.2  mm.).  They  transform  at  %  inch  (14-16  mm.).  Tadpole  reddish  brown, 
23-25  mm. 

Our  evidence  on  transformation  comes  from  two  lots  of  material  (USNM 


HANDBOOK  OF  FROGS  AND  TOADS 

and  AMNH).  The  first  accessions  (USNM  nos.  71030-3)  were  collected  at 
Gainesville,  Fla.,  by  Gerrit  S.  Miller,  Jr.,  and  C.  R.  Ashemeier.  They  measure 
respectively  14, 15.5, 15, 14  mm.  These  specimens  reveal  none  of  the  character- 
istic spots  of  the  adult  except  the  bars  on  the  fore  and  hind  legs.  All  four  have 
an  indication  of  the  dark  band  from  eye  to  shoulder. 

The  other  notes  were  taken  on  19  "Pseudacris  copit"  specimens  collected  by 
T.  Hallman  in  Florida  (AMNH  nos.  15155-8,  15234-47,  15976-7).  My  notes 
designate  them  as  transformed  specimens,  but  evidently  some  of  them  are 
slightly  or  well  past  transformation. 

They  range  from  14-19  mm.  in  length.  Of  two  we  write :  No.  15157 — 18  mm., 
tail  9  mm.,  musculature  light  below,  dark  above.  No.  15158 — 16  mm.,  tail  16 
mm.,  very  spotty  musculature,  dorsal  and  ventral  crests. 

The  measurements  of  all  were:  14, 15.5, 15.5, 16, 16, 16, 16.5, 16.5, 17, 17.5, 18, 
18, 18, 19, 19, 19, 19, 19,  19  mm.  In  1922  Hallinan  also  took  at  Arlington,  Fla., 
a  transformed  specimen  (AMNH  no.  215360).  It  is  just  transformed  and  17 
mm.  long. 

Finally  three  specimens  (Fla.  State  Mus.  no.  47679)  taken  6  miles  southwest 
of  Gainesville,  Fla.,  May  7, 1930,  by  O.  C.  Van  Hyning  and  Norman  W.  Davis 
measure  16, 18,  and  20  mm.,  respectively.  The  first  is  just  transformed  and  the 
others,  though  past,  still  show  signs  of  transformation.  The  transformed  one 
of  16  mm.  has  a  dark  bar  from  eye  to  shoulder  and  a  spot  halfway  from  fore 
limb  to  hind  limb.  The  20-mm.  specimen  has  the  vitta  from  snout  through  eye 
and  onto  sides,  also  oblique  groin  spot  and  oblique  pair  of  spots  on  back  from 
anus  forward. 

We  still  need  detailed  descriptions  of  the  egg  mass,  individual  eggs,  and  tad- 
poles. 

Authorities9  corner: 
F.  Harper,  Ga.,  1937,  pp.  260-272. 
A.  F.  Carr,  Fla.,  1940^  pp.  57-58. 

[Note:  See  comparison  with  P.  strec\en  in  the  account  of  that  species.] 

Strecker's  Ornate  Chorus  Frog,  Texas  Ornate  Chorus  Frog 

Pseudacris  strec\eri  Wright  and  Wright.  Plate  LVIII;  Map  16. 

Range:  Texas,  Oklahoma,  southeastern  Arkansas.  The  Mississippi  record 
may  have  been  a  mistake.  Was  it  P.  brachyphona  or  P.  ornata?  Bragg  (1942) 
records  it  from  19  or  21  counties  in  Oklahoma,  and  we  have  seen  it  from  Bee- 
ville  to  Waco  in  Texas. 

Habitat:  Moist  shady  woods,  grassy  pastures,  among  grain  stalks,  or  in  cot- 
ton fields.  We  found  it  breeding  in  a  semiswampy,  springy  fork  of  a  rocky 
branch  of  Helotes  Creek  in  Texas.  From  our  first  experience  with  it  and  Streck- 
er's  site  (1910)  we  concluded  it  preferred  rocky  ravines  or  deep  gulches,  but 
the  males  are  not  choosy  at  all. 


TREE  FROGS:  HYLIDAE 


273 


Plate  LV1I.  Pseudacris  ornata  (X')-  i-4-  1>late  LVUL  ^udacris  streckfri.  1,3,4,5,6. 

Females.  Males  (XO-  2-  Male  (X%). 


274  HANDBOOK  OF  FROGS  AND  TOADS 

Strecker  writes  (Tex.,  19266,  p.  n) :  "These  frogs  inhabit  all  kinds  of  situa- 
tions. We  found  them  in  moist  shady  woods,  in  weed  clumps  and  among  grain 
stalks  along  the  edges  of  corn,  wheat,  oat,  and  cotton  fields,  among  weeds 
along  the  sides  of  dry,  dusty  roads,  in  grassy  pasture  land  near  lagoons  and 
streams,  and  among  sparse  vegetation  on  the  crests  of  low  rocky  bluffs.  They 
were  lively  and  seemed  perfectly  comfortable  under  the  hot  rays  of  the  sun. 
Several  specimens  flushed  by  us  sprang  into  small  bushes  and  clung  to  their 
branches." 

Size:  Adults,  1-1%  inches.  Males,  25-41  mm.  Females,  32-46  mm.,  possibly 
ripening  as  early  as  27  or  28  mm.  In  Strecker's  collection  there  is  not  much 
evidence  of  ripening  at  28  mm. 

General  appearance:  The  head  is  short  and  wide;  the  body,  short  and 
squatty.  The  fingers  of  the  hand  are  chubby,  forming  a  toadlike  hand.  The 
toes  are  very  slightly  webbed,  the  disks  slight.  The  frog  varies  in  color  from 
pale  drab-gray  or  pale  smoke  gray  to  hazel,  brownish  olive,  or  pea  green. 
Usually  there  are  dorsal  spots  and  frequently  a  triangle  between  the  eyes.  A 
dark  line  extends  from  the  snout  through  the  nostril  and  eye,  over  the  tympa- 
num to  the  arm  insertion;  it  is  sometimes  continued  as  spots  along  the  side. 
Dark  crossbands  are  on  the  legs.  The  groin  and  the  front  and  rear  of  the  femur 
in  males  may  be  yellow  or  green  or  olive-buff.  The  throat  of  the  male  is  dark. 
The  tympanum  is  very  small. 

Color:  Male.  Helotes,  Tex.,  Feb.  16,  i()25.  Back  drab-gray;  triangular  spot 
between  eye  and  spots  on  arms,  legs,  and  back  grayish  olive;  line  on  nostril, 
spot  below  eye,  vitta  back  of  eye,  shoulder  spot,  and  spots  in  a  row  along  the 
side,  black,  and  tympanum  included  in  this  color;  crossbands  on  femur  gray- 
ish olive  behind,  blackish  in  front;  a  black  spot  at  the  insertion  of  arm;  grape 
green  or  dark  olive-buff  in  groin  and  on  front  of  femur;  throat  very  discolored, 
dark  olive-buff  at  its  caudal  edge  shading  forward  into  deep  olive;  under  parts 
white;  underside  of  thighs  slightly  dark  olive-buff  and  buffy  olive.  Iris  vina- 
ceous-fawn  or  light  cinnamon-drab  with  "bronzy  cast."  There  is  a  tendency  for 
a  horizontal  line  of  black  across  the  ins  to  connect  the  nostril  line  with  the 
spot  back  of  the  eye.  Another  specimen  brownish  olive  above  with  no  dorsal 
spots.  Another  one  with  dorsal  spots  of  calla  or  parrot  green.  In  one,  the  spots 
through  nostril,  back  of  eye,  over  shoulder,  and  back  of  shoulder  seal  brown. 
In  this  specimen,  groin,  front  and  rear  of  femur,  and  tibia  are  lemon,  chrome, 
or  aniline  yellow.  Light  color  on  upper  jaw  and  rim  of  lower  jaw  back  to  un- 
der tympanum  is  olive-buff. 

Strecker  (i<)26e)  says,  "The  following  are  the  three  principal  types  of  perma- 
nent marking: 

"A.  Large  and  distinct  black-brown  dorsal  markings  in  regular  pattern. 
(Mostly  males.) 

B.  Distinct  markings  along  both  sides  of  the  dorsal  area,  but  absent  from  the 
middle  of  the  back.  (Mostly  females.) 


TREE  FROGS:  HYLIDAE  275 

C.  Markings  of  the  dorsal  region  broken  up  into  spots  and  flecks.  A  very  dis- 
tinct and  uncommon  type,  confined  to  a  few  female  specimens." 

Of  a  series  of  20  males  and  12  females,  Strecker  (Tex.,  ig26d,  p.  9)  found  a 
range  of  color  thus :  Light  dove  gray,  dark  gray,  light  plumbeous  wood-brown, 
purplish  brown,  chestnut,  bronze-buff,  dull  green,  pea  green,  suffused  with 
green  above.  He  specifies  the  markings  in  each  case. 

Structure:  Tongue  rounded  behind.  Vomerinc  teeth  behind  nares.  Very  lit- 
tle webbing  on  feet.  When  wiping  off  one  male  for  photographing,  I  sensed 
that  it  had  considerable  secretion.  Very  pebbly  or  granular  on  femur  and  ven- 
ter to  pectoral  fold.  Two  metatarsal  tubercles. 

In  preserved  specimens  a  median  plait  on  throat,  or  two  plaits,  or  as  many  as 
six  or  more.  Each  side  of  throat  often  pebbled.  The  male  sometimes,  like  the 
female,  may  have  a  transverse  fold  across  the  pectoral  region.  Some  females 
may  also  have  one  across  the  base  of  the  throat  (in  preserved  specimens). 

E.  D.  Cope's  first  description  of  present  P.  st*cc\en  follows:  "Chorophihts 
oculans  Daudm,  Cope.  This  species  resembles  the  eastern  C.  oculans f  but  some 
specimens  differ  in  the  tuberculate  character  of  the  skin  of  the  superior  sur- 
faces, and  in  the  rudimental  web  of  the  hind  foot.  The  head  is  rather  short,  and 
the  anterior  outline  is  a  narrow  oval.  The  extremity  of  the  muzzle  projects  be- 
yond the  mouth,  and  the  lores  are  slightly  oblique  and  a  little  concave.  The 
nostril  is  but  little  nearer  the  extremity  of  the  muzzle  than  the  orbit.  The  verti- 
cal diameter  of  the  tympanum  a  little  exceeds  the  transverse,  which  is  one-half 
the  long  diameter  of  the  eye-slit.  The  pupil,  as  in  the  other  species  of  this  genus, 
is  horizontal.  The  tongue  is  wide,  discoid,  and  entire  behind.  The  ostia  pharyn- 
gea  are  smaller  than  the  small  choanae.  The  vnmerinc  patches  are  short  and 
transverse;  they  are  entirely  within  the  lines  of  the  inner  borders  of  the  choanae 
and  behind  the  line  of  the  posterior  borders  of  the  same. 

"The  tubercles  of  the  superior  surfaces  are  small  and  rather  closely  placed; 
they  are  largest  on  sides  of  the  back.  There  is  a  faint  arcolntum  of  the  gulnr 
region.  The  limbs  arc  short  and  stout.  The  humerus  is  half  or  more  inclosed  in 
the  skin.  The  palm  reaches  nearly  to  the  end  of  the  muzzle.  The  fingers  are 
short  and  stout,  and  have  neither  dilations  nor  borders.  The  first  is  shorter  than 
the  second,  which  equals  the  fourth.  The  palmar  tubercles  arc  not  distinct.  The 
heel  of  the  appressed  hind  foot  in  thin  specimens  marks  the  middle  of  the  tym- 
panic disc  or  posterior  border  of  orbit,  and  the  end  of  the  muzzle  the  extremity 
of  the  tarsus.  The  hind  foot  beyond  the  tarsus  is  only  as  long  as  the  tibia.  The 
toes  have  no  dilatations,  but  possess  dermal  margins,  and  a  short  but  distinct 
basal  web.  There  is  but  one  solar  tubercle,  a  small  cuneiform  prominence. 
Total  length,  m.,  .035;  of  head,  to  line  of  posterior  borders  of  membramtm 
tympam,  .011;  width  of  head  at  the  latter,  .014;  length  of  hind  leg,  .045;  of 
femur,  .013;  of  hind  foot,  .022;  of  tarsus,  .CXJQ. 

"The  color  above  is  olive-gray,  and  below,  uniform  straw-color.  A  black  band 
passes  from  the  end  of  the  muzzle  on  each  side,  through  the  eye,  and  expand- 


276  HANDBOOK  OF  FROGS  AND  TOADS 

ing  over  the  ear-drum,  terminates  in  front  of  the  humerus.  One  or  two  dark 
spots  above  and  behind  the  axilla  may  unite  to  form  part  of  a  lateral  band. 
There  may  or  may  not  be  blackish  spots  above  the  groin  and  on  the  coccygeal 
region  and  anterior  part  of  the  back.  The  limbs  have  a  few  dark-brown  cross- 
bands;  the  femur  is  yellowish  and  unspotted  behind. 

"There  is  some  difference  between  the  Texas  specimens  and  those  from 
Georgia.  Specimens  from  the  latter  State  are  very  smooth,  and  the  limbs,  espe- 
cially the  feet,  are  slender.  The  heel  reaches  to  the  orbit,  or  at  least  to  the  front 
of  the  tympanic  membrane,  and  the  end  of  the  tarsus  extends  to  or  well  beyond 
the  end  of  the  muzzle.  The  web  and  digital  dermal  borders  are  much  less 
marked.  Two  specimens  were  obtained  by  Mr.  Boll  near  Dallas,  and  three  at 
Helotes  by  Mr.  Marnock.  All  the  latter  have  large  brown  dorsal  spots"  (E.  D. 
Cope,  Tex.,  i88oa,  pp.  27-28). 

Cope  compared  this  Texas  form  with  P.  ornata,  which  he  called  both  C.  ocu- 
lans  and  C.  occidentalis.  Workers  now  have  a  simple  time  in  comparison  with 
our  struggles  when  we  tried  30  to  40  years  ago  to  identify  Chorophtlus  now 
Pscudacns.  From  1900  to  1915  everyone  was  hopelessly  confused  in  terminology 
and  records.  Field  experience,  of  which  we  yet  need  much  more,  was  the  main 
factor  in  clarification. 

Voice:  This  call  is  rather  shrill,  somewhat  like  that  of  the  common  tree  toad. 

"The  calling  habits  of  P.  strecferi  are  distinctive.  The  voice  is  clear  and  bell- 
like  and  has  considerable  carrying  power.  The  breeding  cry  is  a  single  note  but, 
when  heard  in  a  chorus,  the  sound  resembles  the  squeak  of  an  ungreased 
wooden  wheel.  This  effect  results  from  a  lack  of  synchrony  of  individual  calls. 
Males  call  from  various  positions.  I  have  found  them  hanging  to  vegetation  by 
their  hands  in  water  two  feet  deep,  in  vegetation  above  the  water,  and  on  the 
banks  of  pools,  sometimes  as  far  as  five  or  six  feet  from  water.  One  was  found 
sitting  on  a  floating  board  in  a  deep  ditch  calling  lustily.  In  the  more  than  fifty 
cases  lhat  I  have  observed,  P.  strecfon  did  not  stand  'on  tip-toe'  while  calling 
as  does  its  close  relative,  P.  ornata.  .  .  . 

"The  call  of  an  individual  of  this  species  stimulates  other  males  to  call  also. 
I  have  often  approached  a  pool  when  no  males  were  calling.  Within  a  few 
minutes,  when  one  call  was  heard,  a  whole  chorus  would  break  out  and  con- 
tinue lustily  for  ten  minutes  or  more.  Then  all  lapsed  into  silence  again.  After 
an  interval  of  five  to  fifteen  minutes  this  process  would  be  repeated.  This  phe- 
nomenon is  more  prevalent  during  daylight  than  at  night.  Usually  after  dark- 
ness has  fallen,  especially  after  heavy  rains  within  the  breeding  season,  calling 
is  continuous  on  through  the  night'1  (A.  N.  Bragg,  Okla.,  1942^1,  p.  49). 

Breeding:  These  frogs  breed  from  December  to  late  May.  Our  Texas  friends, 
Mr.  Kirn  and  Mr.  and  Mrs.  R.  D.  Quillin,  hold  that  they  are  more  or  less  active 
and  breed  through  the  winter  from  December  to  late  February.  Mr.  Garni  at 
Bocrne,  Tex.,  says,  "They  are  heard  throughout  the  winter,  especially  after 
rains." 


TREE  FROGS:  HYLIDAE  277 

"The  eggs,  like  those  of  Pseudacris  triscnata  (Wied)  are  in  small  bunches, 
usually  from  ten  to  twenty-five.  In  the  ditches  leading  to  Belle  Mead  (Dry 
Pond),  they  are  attached  to  weeds  and  water  grasses,  occasionally  to  twigs  and 
water-logged  branches"  (J.  K.  Strecker,  Jr.,  Tex.,  19266,  p.  n). 

One  egg  mass  in  Strecker's  collection  is  very  large.  There  are  approximately 
200  or  250  eggs  in  one  loose  mass.  Each  egg  is  brown  above  and  cream  below. 
Apparently,  according  to  a  superficial  examination,  there  is  one  envelope, 
which  is  about  2%  or  3  times  the  diameter  of  the  vitellus,  which  is  roughly 
2  mm.  or  less.  Detailed  measurements  of  these  eggs  reveal  the  following: 

The  egg  mass  is  rather  milky  in  its  formaldehyde  preservation  and  has  some 
precipitate  on  it.  There  seems  to  be  only  one  envelope,  though  at  times  appear- 
ances indicate  a  second  inner  one;  but  of  this  we  are  not  certain.  Our  impres- 
sion is  one  envelope,  which  measures  from  3.0  through  3.6, 3.8  to  4.4  or  5.0  mm. 
in  diameter.  The  vitellme  diameter  is  from  1.2-1.8  mm.,  average  1.4,  mode  1.4 
mm. 

"The  tadpoles  are  small,  blackish-brown  creatures  similar  to  those  of  Insert- 
ata,  but  are  rather  shorter  and  more  robust.  .  .  .  About  sixty  days  are  required 
for  them  to  go  through  with  their  complete  transformation  after  leaving  the 
egg"  (J.  K.  Strecker,  Jr.,  Tex.,  19266,  p.  n). 

"The  young  specimens  captured  at  Dry  Pond  near  Waco,  and  at  San  Marcos, 
were  about  twelve  millimeters  in  length,  and  must  have  been  out  of  the  water 
for  several  days,  for  the  dorsal  pattern  was  clear  and  distinct.  When  they  first 
come  to  land,  they  are  dark  gray  with  hardly  any  trace  of  markings"  (same). 

We  have  examined  the  Baylor  material  on  which  the  previous  statement  is 
partially  based.  There  are  frogs,  apparently  just  beyond  transformation,  meas- 
uring from  12  mm.  (Baylor  Mus.  no.  3512)  through  13  (3509)  to  1^.5  (5228) 
mm.  The  body  of  the  largest  tadpole  Strecker  has  is  12  mm.  in  length. 

The  most  precise  information  on  the  eggs  and  tadpoles  comes  from  A.  N. 
Bragg  (Okla.,  1942^  pp.  50-59),  who  has  written: 

"The  eggs  are  attached  to  vegetation  in  masses  of  from  ten  to  about  one 
hundred,  just  under  the  surface  of  the  water.  The  usual  number  in  a  mass  is 
from  twenty  to  fifty  and  adjacent  masses  are  often  loosely  joined  by  thin  sheets 
of  jelly.  .  .  ,  The  egg  complement  is  of  medium  size.  Three  females  produced 
the  following  numbers  of  eggs  in  the  laboratory:  708,  695,  and  401  and  counts 
of  several  sets  of  adjacent  masses  of  eggs  collected  in  the  field  and  presumably 
coming  in  each  case  from  one  female  gave  comparable  values.  More  superficial 
observation  of  about  fifty  sets  offered  evidence  that  the  above  numbers  repre- 
sent fairly  closely  the  usual  yearly  egg  capacity  of  one  female.  .  .  . 

"The  eggs  may  be  described  as  follows:  color,  brownish  gray  at  the  animal 
pole,  the  pigment  extending  downward  to  beyond  the  equator,  then  fading 
rapidly  through  shades  of  lighter  gray  to  cream  or  nearly  pure  white  at  the 
vegetal  pole.  The  animal  pole  is  really  quite  dark  and  the  color  is  only  slightly 
less  intense  at  the  equator.  The  sizes  of  the  eggs  are  nearly  uniform  and  there 


278  HANDBOOK  OF  FROGS  AND  TOADS 

are  no  differences  observable  to  the  naked  eye  as  one  sees  them  en  masse  in 
culture  dishes.  Measurements  of  twenty  live  eggs  by  means  of  an  ocular  mi- 
crometer were  as  follows:  13  measured  1.23  mm.  each;  4, 1.29  mm.;  2, 1.26  mm.; 
and  i,  1.17  mm.  .  .  . 

"The  jelly  is  very  sticky,  almost  absolutely  hyaline,  soft,  and  very  elastic. 
Even  in  clear  water,  tiny  particles  of  soil  and  algae  catch  easily  upon  it  and 
serve  to  protect  the  eggs  from  detection.  In  a  muddy  pool,  the  jelly  collects  a 
complete  coating  of  particles  within  a  half  hour  so  that  its  color  becomes  exactly 
that  of  the  surrounding  water. 

"The  sizes  of  the  individual  masses  of  jelly  of  course  vary  with  the  number 
of  eggs  contained  in  each.  The  portion  of  jelly  between  each  two  adjacent  eggs 
is  also  variable  in  amount  but  the  distance  separating  each  egg  from  the  near- 
est one  adjacent  is  usually  between  5,58  and  7.25  mm.  Staining  of  the  jelly  for 
one  hour  and  forty-five  minutes  in  Schneider's  aceto-carmme  revealed  that 
(i)  only  one  jelly  envelope  occurs  about  each  egg;  (2)  this  envelope  is  finely 
laminated,  the  jelly  apparently  laid  upon  the  egg  in  concentric  spheres,  the  wall 
of  each  of  which  is  thin;  and  (3)  there  is  a  very  thin,  nongelatinous  membrane 
surrounding  a  very  small  cavity  in  which  the  egg  actually  lies." 

Journal  notes:  Feb.  16,  1925,  Hclotes,  Tex.  Anna  says  this  is  a  fat  squatty 
Hylaf  not  a  Pseudacns.  If  is  not  in  a  class  with  Pseudacris  oculans  or  Pseud- 
acris n.  clarion. 

Feb.  16.  Rather  shrill  call.  At  once  it  appealed  as  a  note  new  to  me.  A  note 
somewhat  like  H.  verstcolor.  Soon  found  three  males  calling.  One  was  on  a 
bank,  another  among  vegetation  in  water  3  or  4  inches  deep,  and  another  hang- 
ing to  a  branch  at  the  water.  Put  these  in  a  bag,  and  they  began  to  croak;  had 
three  or  four  diverse  notes  in  the  bag.  These  "bag  frogs"  made  others  croak. 
Caught  five  or  six  in  all.  Do  not  believe  the  species  is  at  its  height.  Air  31°  at 
7:30  P.M.  when  these  were  calling,  water  about  the  same.  This  afternoon  was 
very  humid. 

Feb.  19,  Helotes,  Tex.  During  the  night,  one  or  two  captive  males  called  in 
the  can.  Temperature  close  to  ^2°F.  This  morning  air  in  the  can  is  41°.  Most 
of  the  frogs  are  rather  inactive. 

Feb.  21.  At  9  P.M.  air  70°,  didn't  hear  them.  When  I  returned,  air  about  same; 
heard  one,  then  later  two  at  once,  but  they  soon  stopped. 

Feb.  23.  Tonight  at  8:50  P.M.  when  air  was  «;60F.,  two  ornate  chorus  frogs 
were  calling  at  the  east  end  of  the  "ornate  area,"  and  one  above  it  at  west  end. 
Could  not  find  them.  Water  was  5°  or  10°  warmer. 

Feb.  24.  When  air  was  64°,  8:50  P.M.,  about  three  P.  strec\eri  were  calling. 
They  generally  are  almost  wholly  out  of  the  water.  The  one  I  watched  was 
wholly  so.  Even  when  not  croaking,  the  throat  is  overmuch  swollen.  This  one 
was  quite  brown,  with  hardly  a  dorsal  spot  showing. 

March  24,  Beeville,  Tex.  Air  is  resounding  with  Pseudacris  n.  clartyi.  When 
we  came  to  little  creek,  we  heard  peculiar  note,  went  up  one  road  and  started 


TREE  FROGS:  HYLIDAE 

across  soft  field,  but  gave  it  up,  followed  up  another  road,  and  found  the  one 
peculiar  beast  to  be  a  male  Pseudacris  strecl^eri.  It  was  in  the  middle  of  the  road. 
We  drove  on.  Coming  back,  we  heard  another  in  the  same  place  and  later  a 
lone  voice  near  our  camp.  Is  it  a  land  call  ? 

Feb.  9, 1932.  We  received  13  live  Pseudacris  strec^en  males  from  our  friend 
Albert  J.  Kirn,  of  Somerset,  Tex.  In  a  subsequent  letter  (Feb.  15)  he  spoke  of 
them  as  follows:  "I  sent  by  express  thirteen  of  the  Pseudacris  (ornata)  street^- 
eri:— two  of  them  reddish  (i  cinnamon  and  smooth,  the  other  warty)  two 
green  and  the  remainder  normal  colored.  So  you  will  find  that  at  least  color 
and  smoothness  of  skin  have  nothing  to  do  with  the  stability  of  the  subspecies. 
The  frogs  quit  calling  the  last  night  I  got  them,  or  at  least  I  have  heard  very 
few  since  then.  They  can  be  heard  clearly  for  a  distance  of  more  than  half  mile. 

"I  had  on  hand  three  specimens  of  Pseudacris  in  pickle  (one  small).  I  in- 
cluded them  with  ten  Scaphiopus  hurteni  that  I  mailed  a  few  days  ago." 

Notes  on  these  three  are:  (i)  A  female  35  mm.  long,  practically  devoid  of 
dorsal  and  lateral  spots  or  leg  bars  and  with  no  vitta;  taken  at  Lytle,  Tex., 
Nov.  50,  1931,  by  A.  J.  Kirn.  (2)  A  well-spotted  spent  female,  34  mm.  long. 
Mr.  Kirn's  notes  are:  "Lytle,  Texas,  May  31,  1931.  Dug  into  sandy  soil  about 
i%  inches  to  its  back.  Found  by  our  seeing  the  sand  pushed  up.  Silent  since 
January.  Call  all  winter."  Mr.  Kirn  unhesitatingly  identifies  these  two  as 
Pseudacris  (ornata)  strecfen.  (3)  A  small,  much-spotted,  slim-legged  speci- 
men 26.5  mm.  in  length,  taken  1931  in  Pleasanton,  Tex.,  by  L.  Clayton  Foster 
for  A.  J.  Kirn.  The  latter  merely  identifies  this  specimen  as  Pseudacris.  Were 
it  not  for  the  numerous  dorsal  spots,  we  might  put  it  with  the  Pseudacris  ornata 
of  Florida,  whose  fingers  arc  less  chubby  than  those  of  the  adult  P.  strecl(en9 
but  this  character,  like  the  slender,  prominently  barred  hind  legs,  may  be  an 
indication  of  its  immaturity.  We  provisionally  call  it  P.  strecfyn. 

At  last,  March  24,  1932,  we  have  in  hand  at  the  same  time  live  frogs  of  the 
so-called  form  Pseudacris  occidentals  from  Florida,  and  live  frogs  of  the  so- 
called  form  Pseudacris  ornata  from  Texas.  They  are  different  forms.  There  is 
no  doubt  that  the  form  from  Florida  is  the  Pseudacris  ornata  of  Holbrook  and 
the  form  from  Texas  is  no  longer  P.  ornata.  We  call  it  Pseudacris  ptrecl(cri. 
Several  live  frogs  of  each  form  put  side  by  side  reveal  the  following  from  super- 
ficial examination : 

Pseudacris  ornata  from  Florida  is  the  more  slender  form:  the  snout  is 
pointed;  the  arms  and  legs  are  longer  and  more  slender;  the  fingers  and  toes 
are  long  and  slender,  with  a  mere  trace  of  web  at  base  of  toes.  The  indistinct 
darker  marks  on  the  back  are  bars.  The  nostril  is  equidistant  between  snout 
and  eye;  the  dark  bar  on  the  arm  is  long. 

Pseudacris  strecl(eri  from  Texas  is  a  short,  fat,  squatty  form:  the  snout  is 
shorter  and  broader  at  the  tip;  the  arms  and  legs  are  short  and  broad;  the  fingers 
are  short  and  fat,  giving  the  frog  a  toadlike  hand.  The  dark  vitta  on  the  face 
usually  ends  in  front  of  the  arm  insertion.  The  dark  rim  of  the  upper  jaw  is  con- 


280  HANDBOOK  OF  FROGS  AND  TOADS 

spicuous.  The  light  area  is  broad  at  the  nostril  and  again  back  of  the  eye.  The 
dark  pattern  on  the  back  is  made  of  irregular,  spotlike  bars,  these  frequently 
having  a  fork  at  the  upper  end.  The  marks  on  the  legs  are  only  partial  bars;  the 
spots  are  inclined  to  be  lighter  in  the  center  and  darker  at  the  rim  even  when 
indistinctly  outlined  with  light.  The  bar  between  the  eyes  may  be  a  conspicu- 
ous V.  The  dark  spot  is  at  the  base  of  the  arm  only,  or  slightly  on  the  upper  arm. 

Authorities9  corner: 
J.  K.  Strecker,  Jr.,  Tex.,  1915,  p.  48. 
G.  A.  Moore  and  C.  C.  Rigney,  Okla.,  1942,  p.  78. 

"Dec.  4th.  A  few  Chorus  frogs  heard  late  P.M.  yesterday  and  today  late  P.M. 
Heard  late  last  night  also.  A  few  heard  Nov.  18.  Rainy  at  these  times. 

"Dec.  6th.  Chorus  frogs  calling  last  night  when  I  went  to  bed.  Temperature 
43,  calling  again  tonight  TO  P.M.  45°. 

"12/26.  Chorus  frogs  calling  every  night.  Heard  at  10  A.M.  and  5  P.M.  yester- 
day. Weather  cloudy,  misty. 

"Jan.  19.  Chorus  frogs  heard  calling  at  noon,  and  all  morning  not  so  many 
but  heard.  Weather  clear  and  mild.  Rain  day  before  and  Thursday  night. 

"1/27.  Chorus  frogs,  a  few  calling  now  and  then  all  day  today.  Weather 
cloudy,  mild. 

"2/2.  Many  chorus  frogs  calling  this  evening  and  night.  Weather  milder. 
Temperature  in  evening,  48-53,  misty  weather  for  3  days. 

"2/12.  Chorus  frogs  also  calling  and  today  and  Feb.  6,  also  Feb.  2,  heard 
striped  treefrogs  (P.  n.  clarfoi)  calling  from  water  pools  in  south  part  of  San 
Antonio. 

"2/19.  Foggy  all  day;  milder  after  9  P.M.  yesterday.  Chorus  frogs — a  few 
tonight. 

"2/24.  Weather  in  afternoon  a  little  cooler.  Chorus  frogs  tonight  and  last 
night,  not  many"  (notes,  A.  J.  Kirn,  Somerset,  Tex.,  1944). 

We  named  this  species  after  Dr.  John  K.  Strecker,  Jr.,  for  many  reasons: 

(1)  Dr.  Strecker  was  a  pioneer  herpetological  student  in  Texas  life  histories; 

(2)  he  did  many  personal  favors  for  us  and  for  other  workers;  and  (3)  he  first 
described  this  form  in  his  paper  "Studies  in  North  American  Batrachology : 
Notes  on  the  Robber  Frog  (Lithodytes  latrans  Cope)"  (Trans.  Acad.  Set. 
St.  Louis,  June  14, 1910,  XIX5, 73-82). 

[In  1910  few  knew  Pseudacris  forms,  and  Strecker's  statement  that  Litho- 
dytes latrans  laid  egg  masses  in  water  and  had  tadpoles  in  water  seems,  in 
season  and  description,  to  apply  to  P.  strec\en.  Doubtless  Dr.  Julius  Hurter,  his 
early  herpetological  mentor,  helped  him  identify  the  frog,  for  the  paper  ap- 
peared in  the  St.  Louis  Academy  of  Science  Transactions.  He  (Hurter)  too 
must  have  been  confused  because  in  the  USNM  Hurter  collection  (USNM  no. 
57706)  we  found  P.  strecl^eri  labeled  Lithodytes  latrans.  We  have  examined  all 
the  Baylor  University  material,  and  by  1915,  certainly  by  1926,  Strecker  knew 
the  Pseudacris  well  and  the  fog  over  names  was  clearing.] 


Genus  HYLA  Laurent! 

Maps  18-24 

Anderson  Tree  Frog,  Anderson  Frog,  Anderson's  Hyla,  Anderson  Tree  Toad, 

Green  and  Yellow  Tree  Toad 

Hyla  andersonii  Baird.  Plate  LIX;  Map  19. 

Range:  Central  New  Jersey  (Middlesex  Co.)  to  South  Carolina. 

"About  eight  years  ago  James  Chapin  and  I  found  this  beautiful  Tree  Frog 
at  the  Runyon  Pond  two  miles  south  of  Sayreville,  Middlesex  Co.  [NJ.|.  Since 
that  time  I  have  found  it  at  several  other  localities  in  the  same  region— one  mile 
south  of  Old  Bridge,  about  one  mile  southeast  of  Browntown  and  at  Freneau 
near  Matawan.  These  localities  are  all  in  the  sandy  pine  barren  'island*  north 
of  the  Pine  Barrens  proper.  Thus  this  species  extends  northward  to  within  three 
miles  or  less  of  the  lower  Raritan  River,  its  range  coinciding  at  this  point  with 
that  of  the  Carolina  Chickadee"  (W.  DeW.  Miller,  N.J.,  1916,  p.  68). 

"Considerable  doubt  has  been  cast  on  the  type  locality  record  by  later  work- 
ers. Thus,  the  Anderson  record  'has  always  bothered'  Wright  (1932),  who  re- 
marks: 'Anderson  is  twice  as  far  from  the  coast  as  Southern  Pines,  N.C.  (where 
he  also  took  specimens)  or  Cheraw,  S.C.  is  from  the  coast.  Possibly  the  species 
extends  that  far  inland  along  the  Savannah  River.  .  .  .  Anderson's  tree  frog 
is  a  Lower  Austral  (Austroriparian,  Upper  Coastal,  etc.)  form  and  Anderson, 
S.C.  is  in  Upper  Astral  (Carolinian,  Piedmont,  etc.)  country.' 

"Pickens  (letter,  1938)  points  out  that  at  the  time  the  type  specimen  was  col- 
lected most  of  the  travel  to  the  'Up  Country'  was  inland  from  the  coast.  Miss 
Charlotte  Paine  (later  Mrs.  M.  E.  Daniels),  originally  from  Maine  and  an  in- 
veterate collector,  sent  many  specimens  of  amphibia,  etc.,  to  the  Smithsonian 
Institution  and  she  has  generally  been  credited  with  taking  the  type  of  under- 
sonii.  If  this  was  the  case,  Pickens  goes  on  to  suggest,  she  may  have  picked  up 
the  specimen  in  question  at  some  junction  point,  as  Charleston,  Columbia,  or 
Augusta. 

"Although  I  believe  that  andersonii  may  occur  in  the  Savannah  River  swamp 
well  above  the  Fall  line,  the  question  can  be  satisfactorily  settled  only  by  the 
taking  of  another  specimen  reasonably  close  to  Anderson"  (E.  B.  Chamberlain, 
S.C.,  1939,  p.  21). 

Habitat:  White  cedar  swamps.  We  found  larvae  in  several  pools,  grassy, 
sedgy,  sphagnaceous,  along  a  dense  woody  border  below  one  of  the  lakes  at 
Lakehurst,  N.J. 


282 


HANDBOOK  OF  FROGS  AND  TOADS 


"The  exact  positions  of  ten  individuals  were  located,  of  which  seven  were 
captured.  High-bush  Blueberry  tangles  festooned  with  Green  Briars  made  fur- 
ther investigations  in  this  line  impossible.  The  individuals  are  here  referred  to 
by  number. 


Map  79 

"Numbers  i,  2, 3, 4  and  5  were  in  one  group,  within  50  feet  of  the  main  'pike' 
from  May's  Landing  to  Hammonton.  The  ground  was  covered  quite  evenly 
with  Blueberry  bushes  from  a  foot  to  18  inches  high.  Scattered  Pitch  Pines  up 
to  12  inches  in  diameter  stood  from  10  to  30  feet  apart.  The  ground  was  at 
most  damp,  and  the  only  water  nearby  was  a  shallow  pool  about  30  feet  away 
which  probably  dries  up  in  the  summer.  Near  the  bases  of  the  pines  stood  taller 
Blueberry  bushes,  up  to  three  feet  in  height.  No.  i  was  sitting  on  the  main  stem 


TREE  FROGS:  HYLIDAE  283 

of  a  small  Blueberry  bush,  18  inches  from  the  ground  and  six  inches  from  the 
tip  of  the  bush.  A  Pine  stood  i%  feet  away.  No.  2  was  on  the  leaf  of  a  blueberry 
bush,  2%  feet  from  the  ground  and  1%  feet  from  a  Pine  trunk.  No.  3  was  on  a 
little  twiglet  growing  out  from  the  trunk  of  a  pine  3%  feet  from  the  ground. 
No.  4  was  on  the  ground  at  the  base  of  a  Pine.  No.  5  was  one  foot  from  the 
ground,  where  the  twig  of  a  Blueberry  bush  lay  against  the  trunk  of  a  Pine. 
All  of  the  specimens  in  this  group  showed  a  strong  preference  for  the  near 
vicinity  of  a  Pine. 

"Nos.  6,  7,  and  8  were  in  a  thicket  of  small  Red  Maples  and  high  Blueberry 
bushes  in  a  creek  'bottom/  No.  6  was  on  the  main  stem  of  a  Blueberry  bush 
about  four  feet  from  the  ground.  No.  7  was  similarly  located.  No.  8  was  about 
six  feet  from  the  ground  in  a  small  Red  Maple. 

"Nos.  9  and  10  were  in  a  thick  tangle  of  high  Blueberry  bushes  and  Smilax. 
Both  were  near  the  tops  of  Blueberry  bushes  at  least  nine  feet  from  the  ground. 
For  fifty  feet  around  none  of  the  vegetation  was  any  lower,  so  it  seems  that  these 
individuals  climbed  higher  than  is  usual  for  the  species  in  order  to  be  out  in 
the  open. 

"Not  all  of  the  individuals  were  as  tame  as  is  generally  noted  for  andersonii. 
A  number  of  individuals  would  not  continue  singing  when  the  observer  turned 
the  light  on  them  or  approached  nearer  than  fifteen  or  twenty  feet,  and  so  could 
not  be  located.  A  silent  andersonii  in  a  thick  tangle  of  Blueberry  bushes  could 
give  points  on  hiding  to  a  very  small  needle  in  a  very  large  haystack.  No  fe- 
males were  taken"  (A.  B.  Klots,  N.J.,  1930,  pp.  108-111). 

Size:  Adults,  1^-1%  inches.  Males,  30-41  mm.  Females,  38-47  mm. 

General  appearance:  This  is  a  small  green  tree  frog.  The  light-bordered, 
plum-colored  band  along  the  side  of  the  body,  with  its  yellow  spots  below, 
gives  this  beautiful  little  frog  its  distinctive  character.  This  band  marks  its 
green  dorsal  color  very  sharply  from  its  white  ventral  parts.  It  has  orange  in 
axilla  and  groin  and  on  the  rear  of  the  femur.  The  throat  of  the  female  has  a 
white-bordered  green  patch  on  either  side.  In  its  stout  body  it  differs  from  the 
more  slender  and  larger  Hyla  cinerea. 

Color:  Male.  Upper  parts  (dorsum,  upper  lip,  angle  of  mouth,  and  dorsal 
surfaces  of  limbs)  cress  green  to  a  light  cress  green,  on  sides  lighter  to  deep 
chrysolite  green.  Stripe  along  side,  behind  vent,  along  limbs  and  upper  jaw 
cartridge  buff,  ivory  yellow,  marguerite  yellow,  or  seafoam  yellow.  Area  back 
of  eyes,  along  side  to  vent  vinaceous-drab  or  purple-drab,  becoming  over  tym- 
panum dark  purplish  drab.  A  little  of  the  same  color  scattered  on  throat.  Tops 
of  forefeet  and  hind  feet  deep  brownish  drab  except  for  first  two  digits,  which 
have  the  cadmium  yellow  spots  on  the  top.  Fore  part  of  underside  of  antebra- 
chium  and  under  part  of  brachium  with  cadmium  yellow,  deep  chrome,  or 
orange  spots;  the  same  color  also  in  axilla,  groin  and  most  of  fore  and  hind 
parts  of  femur,  whole  undersides  of  tibia,  and  inner  side  of  foot.  All  these 
orange  or  deep  chrome  spots  are  on  a  raw  sienna  or  mars  yellow  background. 


HANDBOOK  OF  FROGS  AND  TOADS 


Plate  LIX.  Hyla  andersonh  (Xrl/0-  i~3 
Males. 


Iris  more  or  less  vinaceous-drab  or 
purple-drab  with  vinaceous-tawny 
spots.  Under  parts  white  except  for 
dark  purplish  drab  of  throat. 

Female.  Throat  white,  grayish,  or 
with  slight  purplish  drab  tinge.  Green 
patch  below  angle  of  mouth  usually 
white-edged,  this  usually  absent  in 
males. 

Structure:  Form  stout,  head  broad 
and  flat;  skin  smooth;  posterior  sur- 
face of  femur  spotted ;  vocal  sacs  sub- 
gular, 

Voice:  This  frog  calls  aquac\- 
aquack^aquacl{t  many  times,  perhaps 
twenty,  at  infrequent  and  irregular 
intervals. 

Dr.  C.  C.  Abbott  (N J.,  1890,  p.  189) 
says,  "The  andersonii  utters  a  single 
note,  better  described  by  the  syllable 
lkeck/  which  it  usually  repeats  three 
or  four  times.  It  is  not  a  frog-like  note 
at  all,  but  much  resembles  the  call  of 
the  Virginia  rail  (Rallusvirginianus) ." 

"June  16-28, 1928.  To  the  writer  the 
call  seemed  a  nasal  'quack/  almost 
verging  on  a  'quank'  but  without  the 
strong  V  sound  of  the  latter.  The  call 
was  never  disyllabic. 

"The  note  is  repeated  at  about  half- 
second  intervals  for  sometimes  fully 
30  seconds.  When  the  frogs  are  in  full 
song  an  interval  of  about  two  minutes 
intervenes  between  outbursts.  We  had 
no  difficulty  in  starting  the  frogs  call- 
ing again  at  distances  of  from  fifty  to 
three  feet,  after  they  had  been  silent 
for  a  minute  or  so.  One  individual 


was  recorded  as  having  called  74  times  in  one  period  of  song. 

"The  frogs  definitely  associate  together  for  singing,  whether  because  of  the 
presence  of  females  or  for  companionship.  The  latter  probably  plays  a  consid- 
erable share  in  the  performance,  as  is  evidenced  by  the  quick  response  to  an 
imitation  of  the  call.  Five  such  singing  groups  were  definitely  located.  Of  these 


TREE  FROGS:  HYLIDAE  285 

the  first  contained  seven  individuals,  the  second  contained  three,  the  third  con- 
tained eight,  the  fourth  contained  three  and  the  fifth,  which  was  just  across  an 
uncrossable  creek,  contained  at  least  six.  Only  once  was  a  single  individual 
noted  in  song  alone,  and  that  was  a  frog  which  called  three  times  in  a  spot  a 
half-mile  distant  from  any  others  and  was  never  heard  from  again. 

"The  locations  of  the  groups  were  fixed,  and  during  our  stay  did  not  change 
a  particle.  Night  after  night  a  group  would  be  in  exactly  the  same  area,  though 
the  individuals  composing  it  shifted  position  a  bit. 

"The  time  of  singing  was  remarkably  constant.  On  every  night  but  one 
[June  16-28,  1928]  the  chorus  started  between  ten  and  fifteen  minutes  before 
sundown.  On  the  one  exception,  a  clear  dry  night  with  a  bright  moon,  the  first 
songs  were  not  heard  until  twenty  minutes  after  sundown. 

"The  carrying  power  of  the  song  was  excellent.  A  chorus  was  plainly  heard 
as  an  entity  over  800  paces  away,  with  two  patches  of  woods  and  a  brushy 
swamp  intervening.  The  wind  was  negligible.  Individual  voices  were  distin- 
guishable 754  paces  away  down  a  straight  road,  with  a  light  wind  blowing 
from  the  observers  toward  the  frogs"  (A.  B.  Klots,  N.J.,  1930,  pp.  108-111). 

"At  dusk  we  have  usually  taken  our  supper,  and  then  waited  for  darkness 
to  come  on  and  for  the  Hylas  to  begin  to  sing;  we  have  had  good  luck  taking 
Hylas  by  the  following  method:  One  of  us  with  an  electric  flashlight  would 
start  for  the  nearest  singing  Hyla,  while  the  other  waited  some  distance  away. 
As  soon  as  the  Hyla  stopped  singing,  the  person  who  was  not  trying  to  ap- 
proach would  imitate  the  call  of  the  frog,  and  this  would  start  it  singing  again 
vigorously,  and  while  it  was  singing  the  collector  bearing  the  light  would  ap- 
proach as  quickly  as  possible,  standing  still  as  soon  as  the  singing  ceased.  This 
process  was  kept  up  until  finally  the  light  flashed  on  the  vibrating  white  throat 
of  the  singing  Hyla,  and  its  capture  then  became  a  perfectly  simple  matter,  as 
they  stared  stupidly  at  the  brilliant  light.  .  .  .  This  year,  however,  we  did  not 
get  down  to  Lakehurst  until  the  8th  of  July,  when  we  found  the  Hylas  singing 
in  goodly  numbers  in  the  white  cedars  about  the  lake.  After  capturing  a  num- 
ber of  singing  males  (I  had  never  taken  a  female  before),  my  light  flashed  by 
merest  chance  upon  a  pair  of  Hylas  sitting  well  up  in  a  pine  tree,  in  embrace. 
This,  and  another  taken  in  a  similar  situation,  were  the  only  females  secured, 
although  we  took  several  males  from  the  low  oak  shrub  about  a  small  fresh 
water  pool  in  the  pine  barrens"  (T.  Barbour,  N.J.,  1916,  pp.  6-7). 

Breeding:  This  species  breeds  from  May  i  to  July  20.  The  eggs  are  single, 
attached  to  sphagnum  or  on  the  bottom.  The  egg  is  /4o-M«  inch  (1.2-1.4 
mm.),  the  inner  envelope  Ki>  inch  (1.9-2.0  mm.),  the  outer  H-%  inch  (3.5- 
4.0  mm.).  The  olive  tadpole  is  small,  i%  inches  (35  mm.),  its  tail  medium 
long  with  tip  acuminate.  The  tooth  ridges  are  %.  After  50  to  75  days,  the  tad- 
poles transform  from  the  end  of  June  to  September  i  at  %«-%  inch  (11-15 
mm.). 

Noble  and  Noble  (N.J.,  1923)  say  that  the  eggs  are  single,  scattered  among 


286  HANDBOOK  OF  FROGS  AND  TOADS 

waterweeds,  attached  to  sphagnum,  or  free  on  the  bottom,  in  small,  nonstag- 
nant  pools  or  in  slow-moving  streams  of  the  pine  barrens.  Color,  dark  brown 
and  creamy  white.  Egg  1.2-1.4  mm-  Inner  envelope  1.9-2  mm.  Outer  envelope 
3.5-4.0  mm.  Complement  800-1000. 

"When  the  tail  is  nearly  absorbed  and  they  leave  the  water,  they  are  about 
25  mm.  long  and  of  a  dull  olive  green.  They  grow  lighter,  that  is,  brighter  green 
in  hue  with  the  disappearance  of  the  tail,  until  the  little  frogs,  which  in  length 
of  body  are  15  mm.,  resemble  the  mature  individuals.  The  white  that  margins 
the  green  of  the  back  and  extremities  is  not  so  conspicuous  as  in  the  adults,  and 
the  saffron  of  the  underparts  is  wanting  in  those  that  I  have  examined"  ( W.  T. 
Davis,  N.J.,  1907,  p.  50). 

Journal  notes:  On  June  8,  1922,  on  our  way  to  Okefinokee  Swamp,  we 
camped  in  the  evening  on  the  north  side  of  Everett's  Pond,  N.C.,  near  the 
S.  Car.-N.  Car.  line.  The  instant  it  became  dark  we  heard  plenty  of  Acns  gryl- 
lus,  Bufo  jowleri,  and  Rana  catesbeiana,  a  few  Rana  clamitans  and  Hyla  cinerea 
in  chorus,  and  Hyla  versicolor.  .  .  .  Later  heard  Hyla  andcrsonii.  These  H. 
andcrsomi  were  in  a  stream  or  branch  with  sweet  gum,  tangle  of  oaks,  bamboo 
briars  (Smilax),  Magnolia  glauca,  maples,  black  gum.  Later  in  the  evening 
Miles  Pirnie  heard  one  opposite  our  camp.  All  the  frogs  heard  were  in  or  near 
the  lake.  Harper  speaks  of  the  calling  of  these  Hylas  thus:  "I  heard  the  note 
of  H.  andersonn  at  a  distance  of  200  yards  and  suspected  almost  at  once  what 
it  was.  Trailed  it  and  after  a  long  wait  located  it  in  some  tall  bushes  in  the  edge 
of  a  branch  swamp.  Its  note  bears  a  general  resemblance  to  that  of  H.  cinerea 
but  goes  about  twice  as  rapidly,  is  about  half  as  loud  and  sounds  more  like  quak, 
quak  than  quonk.  It  carries  tairly  well  at  200  yards  and  about  300  yards  would 
probably  be  the  limit.  .  .  .  Did  not  see  it  croak.  Its  periods  are  infrequent  and 
irregular,  2  minutes'  interval.  Perhaps  15  or  20  or  25  notes  given  in  one  period." 

On  June  28,  1929,  we  heard  these  Hyla  andersonn  in  the  wooded  edge  of 
Mr.  P.  H.  Emilie's  lake  at  Lakehurst,  NJ.  The  males  were  among  magnolias, 
maples,  huckleberries,  azaleas,  Ilex  glabrat  Viburnum. 

Authorities'  corner: 

W.  T.  Davis,  N.J.,  1905,  p.  795.  C.  S.  Brimley,  N.C.,  1940,  p.  22. 

W.  T.  Davis,  N.J.,  1907,  p.  49.  L.  R,  Aronson,  N.J.,  1943,  pp.  246-249. 

G.  K.  and  R.  C.  Noble,  N.J.,  1923,  pp. 

419  ff. 

H.  andersonii  is  a  handsome  species  and  it  has  been  superbly  illustrated. 
From  the  time  of  Miss  Dickerson's  beautifully  colored  plate  in  her  fine  Frog 
Boot^  (Chapin's  specimens  1906-1908),  and  L.  A.  Fuertes'  painting  of  them  to 
the  days  of  our  private  preparatory  tips  to  Dr.  Noble's  well-illustrated,  first 
(1923)  outdoor  life  history  of  H.  andersonii,  we  have  been  interested  in  this 
form.  Several  students,  Mr.  and  Mrs.  A.  B.  Klots,  W.  H.  Caulwell,  and  others, 
have  brought  us  specimens,  particularly  in  the  period  since  1923. 


TREE  FROGS:  HYLIDAE  287 

Canyon  Tree  Frog,  Canyon  Tree  Toad,  Desert  Tree  Toad,  Cope's  Hyla, 
Arizona  Tree  Frog,  Sonoran  Tree  Toad 

Hyla  arenicolor  Cope.  Plates  LX,  LXI;  Map  20. 

Range:  Southwestern  United  States :  western  Texas  through  extreme  south- 
ern Colorado  and  Utah  to  the  region  of  the  Colorado  River  in  Nevada,  down  it 
to  Lower  California  and  up  southern  California!!  coast  to  Ventura  County.  In 
Mexico  to  Guadajuata,  Guadalajaca,  and  Toluca. 

Habitat:  Rocky  canyons.  From  Fern  Canyon  in  Ft.  Davis  Mts.,  Tex.,  to 
Tahquitz  Canyon,  Calif.  Invariably  we  found  them  in  rocky,  clear  streams. 
They  have  been  reported  by  others  from  mountain  springs,  irrigation  ditches, 
and  "rapidly  flowing  streams." 

"Throughout  this  region  [Southern  California],  its  habitat  appears  to  be 
confined  to  streams  and  mountain  springs  between  1000  and  5000  feet  eleva- 
tion. Here  the  writer  has  found  it  associated  with  such  trees  as  Alnus  rhombi- 
jolta,  Platanus  racemosa,  and  Acer  macrophyllum  and  in  this  state  at  least  it 
may  be  regarded  as  an  inhabitant  of  canyons  within  the  upper  sonoran  zone. 

"It  is  apparently  more  strictly  aquatic  than  the  smaller  Hyla  regilla  Baird 
and  Girard,  whose  range  in  Southern  California  is,  in  part,  co-extensive  with  it. 
The  former  species  has  never  been  found  far  away  from  the  vicinity  of  water, 
while  the  latter  has  often  been  seen  under  vegetation  a  considerable  distance 
from  it"  (C.  H.  Richardson,  Jr.,  Gen.,  1912,  p.  607). 

"Occupies  chiefly  the  Upper  Sonoran  life-zone,  extending  locally  into  Lower 
Sonoran.  Lives  on  boulders  and  exposed  rock  faces  close  to  canon  streams" 
(J.  Grinnell  and  C.  L.  Camp,  Calif.,  1917,  p.  145). 

"Fairly  common  around  shady  moist  places  in  the  Canyon  and  occasionally 
up  to  South  Rim.  Lower  Sonoran  and  Upper  Sonoran  Zones"  (E.  D.  McKee 
and  C.  M.  Bogert,  Ariz.,  1934,  p.  178). 

"Common  in  the  pine-fir  and  chaparral-woodlands  zones  but  uncommon  in 
the  semidcsert.  Calling  individuals  and  clasping  pairs  were  common  in  water 
at  a  dam  in  the  pine-fir  forest  on  May  19, 1936.  After  a  mating  season  adults  are 
found  occasionally  about  buildings"  (E.  S.  Little,  Jr.,  Ariz.,  1940,  p.  262). 

"This  species  is  confined  to  the  High  Plateaus-Colorado  River  region  of 
Utah.  They  are  common  in  Zion  National  Park  and  St.  George.  .  .  .  This 
species  is  commonly  found  on  the  rocks  and  banks  of  the  small  streams  and 
pools  of  the  Colorado  River  drainage  of  this  state.  It  is  seldom  if  ever  found 
farther  than  a  few  feet  removed  from  running  water"  (V.  M.  Tanner,  Utah, 
1931,  pp.  186-187). 

Size:  Adults,  i%-2%  inches.  Males,  29-5?  mm.  Females,  30-54  mm. 

General  appearance:  This  tree  toad  averages  smaller  in  size  and  duller  in 
color  than  our  common  tree  toad  (Hyla  versicolor).  When  sitting  quietly  it 
(a  specimen  from  Grand  Canyon,  Ariz.)  looks  surprisingly  like  the  little 
spotted  canyon  toad  (Bufo  punctatus),  but  when  it  climbs,  its  conspicuously 


288  HANDBOOK  OF  FROGS  AND  TOADS 

long  legs  and  large  disks  on  fingers  and  toes  immediately  show  it  to  be  a  tree 
toad.  It  easily  clings  to  a  vertical  surface  and  climbs  up  even  a  sheet  of  glass. 
The  back  is  brown  or  grayish  in  color  with  dark  dots  scattered  over  it.  The  legs 
are  barred  with  dark  areas.  There  is  much  yellow  or  orange  on  the  rear  of  the 
legs,  in  the  groin,  and  in  the  axilla.  It  is  light  beneath.  The  skin  is  slightly 
rough,  becoming  more  so  as  it  becomes  dry.  Unlike  the  Pacific  tree  frog  (Hyla 
regilla),  this  species  has  no  stripe  through  the  eye  and  along  the  side  of  the 
body.  The  skin  is  roughened  in  H.  aremcolor,  smooth  in  H.  regilla.  This  frog 
has  slight  webs  between  the  fingers. 

"Size  small,  head-and-body  length  50  mm.  (2  inches)  or  less;  fingers  and 
toes  with  expanded  adhesive  disks;  no  webs  between  fingers;  dorsal  skin 
rough-surfaced,  with  many  small  papillae;  side  of  head  concolor  with  rest  of 
head  and  body;  coloration  light  or  dark  grny.  .  .  .  Distinguished  from  other 
Salientia  of  California  by  expanded  disks  on  fingers  and  toes  and  by  small  size 
of  adults;  from  Hyla  regilla  by  average  larger  size  (especially  of  females),  by 
larger  discs  on  digits,  by  rougher  skin  on  dorsal  surface  of  body,  and  by  lack 
of  dark  stripe  on  side  of  head  through  eye"  (T.  I.  Storer,  Calif.,  1925,  p.  204). 

Color:  Scntenac  Canyon,  Calif.,  from  L.  M.  Klauber,  March  31,  1928.  Male. 
Upper  parts  olive-huff,  drab,  light  grayish  olive  or  deep  olive-buff.  Snout  ahead 
of  eyes  avcllancous  or  wood  brown,  some  of  larger  warty  spots  with  touch  of 
same  color  or  pale  cinnamon-pink  to  vinaceous-cinnamon.  Bar  between  eyes, 
line  in  front  of  eye  and  from  eye  through  ear,  four  or  five  larger  spots  on  back, 
three  crossbars  on  femur,  four  on  tibia,  four  on  hind  foot,  and  two  on  forearm 
are  from  deep  olive-buff  to  citrine-drab.  These  spots  are  outlined  with  broken 
black  and  filled  in  with  body  color  giving  the  deep  olive-buff  or  citrine-drab 
combination.  Interspaces  between  bars  on  legs  and  arms  vinaceous-buff  or  avel- 
laneous.  Groin,  olive-ocher,  yellow  ochcr,  lemon  chrome,  becoming  aniline 
yellow  or  raw  sienna  on  front  and  rear  of  femur.  Dorsal  color  on  femur  very 
narrow.  Dorsal  color  extends  around  vent;  with  tilleul  buff  or  pale  olive-buff 
on  each  tubercle.  Ventral  view  of  legs  and  arms  vinaceous-fawn  or  light  russet- 
vinaccous  or  light  grayish  vmaceous.  Venter  white  or  cartridge  buff.  Vocal  sac 
when  inflated  light  buff  except  near  chin  where  spot  below  eye  is  light  pinkish 
cinnamon  or  sulphine  yellow.  Iris  with  benzo  brown,  cinnamon-drab,  or 
brownish  drab  band  in  front  and  back  of  pupil,  this  finely  dotted  with  olive- 
buff;  iris  below  and  above  pupil  largely  olive-buff  with  wavy,  more  or  less 
vertical  lines  of  brownish  drab;  pupil  rim  above  dull  green-yellow  or  chalced- 
ony yellow,  light  below;  pupil  rim  below  with  vertical  slit  of  black.  Another 
male  has  back  pale  olive-buff  or  pale  drab  gray. 

Another  male.  Dorsal  color  pale  olive-gray  or  pale  smoke  gray  to  light 
grayish  olive  with  light  vinaceous-cinnamon  on  snout  and  forelegs.  Belly  not 
white  but  dark  olive-buff. 

Female.  Underparts  white  or  pale  olive-buff,  middle  of  belly  or  rear  belly 
olive-buff.  Groin  with  very  little  lemon  chrome. 


TREE  FROGS:  HYLIDAE 


289 


Plate  LX.  Hyla  arenicolor.  (XO-  Part 
i.  1-5.  From  Grand  Canyon,  Ariz.  6,7. 
Transformed  and  tadpole,  from  Fern  Can- 
yon, Davis  Mts.,  Tex. 


Plate  LXL  Hyla  arcnicolor.  (XO-  Part 
2.  From  California. 


29o  HANDBOOK  OF  FROGS  AND  TOADS 

Another  female.  Fern  Canyon,  Tex.,  July  9,  1925.  Spots  on  back,  body, 
legs,  head  mouse  gray  to  dark  olive-gray;  back  deep  grayish  or  olive  becoming 
pale  smoke  gray  on  the  sides  next  to  belly  color,  on  labial  margin,  snout,  and 
.vent  region.  In  axilla,  side  of  lower  belly,  front  and  rear  of  thigh,  rear  of  tibia 
and  inner  face  of  foot,  rear  of  humerus,  somewhat  on  inner  face  of  forearm 
olive-ocher  or  aniline  yellow  or  sulphme  yellow,  becoming  on  the  edges  green- 
yellow.  Breast  white.  Throat  white  spotted  with  mouse  gray  or  dark  olive  gray, 
especially  around  edge  of  lower  jaw.  Tympanum  with  some  orange  rufous  or 
tawny  in  lower  half. 

"The  color  phases  run  from  light  gray  to  brownish  black,  with  numerous 
roundish  spots  irregularly  distributed  and  not  forming  well-marked  patterns, 
except  on  the  hind  legs,  which  appear  banded.  In  the  light  and  dark  color 
phases  the  spots  become  obscure"  (G.  P.  Engelhardt,  Utah,  1918,  p.  79). 

Structure:  Long  arms  and  legs  with  large  disks  on  tips  of  fingers  and  toes; 
slight  webs  between  the  fingers;  skin  somewhat  roughened;  more  or  less  uni- 
form in  color;  tympanum  small;  prominent  fold  across  breast;  prominent  fold 
on  tarsus. 

A.  A.  W.  has  long  held  it  likely  that  there  is  more  than  one  race  of  Hyla 
arenicolor.  The  following  observations  are  some  of  her  notations:  In  general, 
the  frogs  from  California  have  longer  legs  and  a  black  bar  across  the  head,  a 
lighter  gray  background  with  inky  spots,  and  a  more  pebbly  skin.  The  frogs 
from  Grand  Canyon,  Ariz.,  are  smoother,  have  shorter  legs,  arc  reddish  in 
color,  spotted  rather  indistinctly,  and  bear  no  marked  triangle  on  the  head. 
Those  from  Rainbow  Bridge  collected  by  Eaton  are  like  the  Canyon  ones. 
Those  from  the  White  Mountains  are  rather  pebbly,  and  the  one  from  New 
Mexico  is  pebbly.  The  Canyon  frogs  had  larger  disks  on  the  toes,  larger  eyes, 
and  less  distinct  spots.  In  life  they  sat  with  body  parallel  to  finger  when  held, 
in  contrast  with  the  more  erect,  toadlike  posture  of  the  California  frogs. 

"Hyla  affinis,  Baird. — Body  rough.  Tympanum  two-thirds  the  size  of  eye. 
Tibia  not  quite  half  the  length  of  the  body,  but  reaching  more  than  halfway 
from  anus  to  center  of  eyes.  Color  ash  gray  or  green,  with  numerous  rounded 
dorsal  blotches.  Three  transverse  bands  on  each  thigh  and  leg.  No  vermicula- 
tion  on  anterior  and  posterior  faces  of  hind  legs,  nor  on  lower  part  of  sides.  A 
light  spot  under  the  eye.  Web  of  hand  extending  only  to  the  third  joint  of  the 
second  finger.  Arm  from  elbow  less  than  tibia,  but  longer  than  hind  foot. 
About  i%  inches  long.  Hab.  Northern  Sonora"  (S.  F.  Baird,  Gen.,  1854,  p.  61). 

"General  aspect  of  H.  versicolor,  having  the  same  squat  appearance,  the 
granulated  skin  above  and  below,  the  ash-color  back  with  darker  mottlings, 
the  white  spot  under  the  eye,  etc.  The  most  conspicuous  distinctive  features  are 
the  absence  of  webs  ot  the  fingers,  the  greater  length  of  the  hind  legs,  and  the 
blotches  on  the  back  being  in  round  spots,  not  cuneiform.  The  legs  with  three 
bars  not  two,  and  without  the  reticulate  markings  behind  and  below"  (E.  D. 
Cope,  Gen.,  1889,  p.  369). 


TREE  FROGS:  HYLIDAE  291 

Voice:  This  call  has  been  described  as  the  "quack  of  a  duck"  by  Storer,  and 
as  "the  bleat  of  a  goat"  by  Duges.  To  us  the  note  sounded  lower,  much  weaker, 
and  less  blatant  than  that  of  Hyla  versicolor  and  not  so  persistent.  "It  was  like 
a  hoarse  sheep's  'Ba-a-a'  when  they  were  in  the  open,  but  more  like  a  roar  when 
they  were  under  the  rocks"  (T.  H.  Eaton,  Ariz.,  1935^  p.  7). 

"The  frogs  are  most  vociferous  in  the  late  afternoon  and  rarely  are  heard  at 
night"  (G.  P.  Engelhardt,  Utah,  1918,  p.  79). 

"The  song  of  this  tree  toad  is  one  of  the  most  melodious  of  all  our  species" 
(V.  M.  Tanner,  Utah,  1931,  p.  187). 

"The  note  differs  markedly  from  that  of  our  common  western  treetoad 
(Hyla  regilla)  in  being  lower  in  pitch,  somewhat  weaker  in  volume,  and  with- 
out any  tendency  toward  a  two-syllable  sound  such  as  is  heard  often  from 
regilla.  On  two  occasions,  in  San  Diego  County,  the  two  species  were  heard 
side  by  side  and  there  was  not  the  slightest  difficulty  in  distinguishing  them. 
Two  separate  individuals  of  arenicolor  which  were  timed  by  the  watch  were 
croaking  at  one-second  intervals.  With  all  the  males  noted  on  Tahquitz  Creek 
(fully  25  were  located)  only  one  female  was  seen.  The  chorus  began  about 
4  P.M.,  soon  after  the  sun  had  disappeared  behind  the  San  Jacinto  Range,  was 
strongest  just  after  dark,  from  about  7:30  to  8:30  P.M.,  and  continued  on  into 
the  night  at  least  until  2  A.M.  On  other  occasions  males  of  Hyla  arenicolor  were 
heard  croaking  up  to  about  6  A.M.,  but  only  a  few  notes  were  given  as  late  as 
that  hour  in  the  morning.  The  duration  of  the  'song'  season  is  unknown;  on 
the  morning  of  June  21, 1919,  Canon  Tree-toads  were  heard  croaking  in  canons 
near  the  Mount  Wilson  trail"  (T.  I.  Storer,  Calif.,  1925,  p.  208). 

"Tree  toads  of  this  species  were  found  in  White  House  Canyon,  in  the  Santa 
Rita  Mountains,  August  3.  Males  were  heard  calling  just  before  dark  from 
branches  of  oak  trees,  about  fifteen  feet  from  the  ground.  The  call  was  an  even, 
strong  trill"  (F.  W.  King,  Ariz.,  1932,  p.  176). 

Breeding:  This  species  breeds  from  March  i  to  July  i.  The  eggs  are  single, 
floating  near  the  surface  or  on  the  bottom  of  the  pool  usually  attached  to  leaves 
(Atsatt  and  Storer).  The  tadpole  is  medium  in  size,  2  inches  (50  mm.)  long, 
dark  olive  in  color  with  some  tail  crests  suffused  with  reddish,  orange-pink  to 
coral  red.  The  tooth  ridges  are  %.  After  a  period  of  40  to  75  days,  the  tadpoles 
transform  from  June  i  to  August  15,  at  %  inch  (14-17  mm.).  We  have  seen 
mature  tadpoles  collected  in  April  from  Fern  Canyon,  Ft.  Davis  Mts.,  Tex., 
and  neither  in  1917  nor  in  1925  did  we  hear  the  choruses  after  July  i,  and  rarely 
single  voices.  C.  H.  Richardson,  Jr.  (Gen.,  1912,  p.  607),  indicates  "that  the 
breeding  season  .  .  .  extends  from  late  spring  until  fall."  The  extension  of 
the  breeding  season  to  fall  is  apparently  based  on  one  female  taken  Sept.  4, 
1906,  and  containing  large  eggs. 

"White  House  Canyon,  in  the  Santa  Rita  Mountains.  .  .  .  Tadpoles  of 
H.  arenicolor  were  found  in  pools  nearby  with  some  of  the  larvae  transform- 
ing at  18  mm.  body  length"  (F.  W.  King,  Ariz.,  1932,  p.  176). 


292  HANDBOOK  OF  FROGS  AND  TOADS 

"I  have  found  it  to  be  fairly  common  at  the  Water  Cress  Spring  in  St.  George 
in  February  and  on  August  28, 1919 1  found  newly  hatched  as  well  as  matured 
tadpoles  in  the  'Stadium  pool'  in  Zion  National  Park.  The  tadpoles  are  black 
when  small  becoming  grayish  when  older"  (V.  M.  Tanner,  Utah,  1931,  pp.  186- 
187). 

"Ova  were  observed  in  the  form  of  small  clusters  deposited  along  the  margin 
of  pools.  The  tadpoles,  at  first  black,  later  become  mottled  gray  when  they  re- 
sort to  deeper  water"  (G.  P.  Engelhardt,  Utah,  1918,  p.  79). 

Journal  notes:  July  16,  1917,  Pinaleno  Mts.,  Ariz.  On  the  rocks  over  the  wa- 
ter, Anna  put  her  hand  accidentally  on  a  Hyla  aremcolor  and  Ray  Shannon 
found  another  on  a  tree  trunk  about  4  feet  up.  Paul  Needham  found  another 
on  a  boulder  just  above  the  water.  They  do  not  seem  at  home  in  the  swifter 
water. 

July  5,  1925,  Fern  Canyon  (near  Alpine,  Tex.).  In  the  ravine  just  after  a 
drenching  ram,  we  found  four  transformed  Hyla  aremcolor  on  the  boulders 
some  25  or  more  feet  above  the  level  of  the  creek.  The  creek  was  full  of  them 
before  the  rain.  Today,  July  6,  the  creek  is  down  and  clear.  Near  the  falls  we 
often  found  one  or  two  Hyla  aremcolor  tadpoles  amongst  the  boulders  in  shal- 
low water  or  swimming  at  the  surface.  We  secured  today  only  one  mature 
tadpole.  The  rest  were  small  tadpoles.  At  the  pool  below  the  falls  were  many 
of  them.  .  .  .  Also  some  above  the  falls. 

July  7,  Ranger  Canyon.  About  6:45-7  P.M.,  we  arrived  at  the  big  pool  or 
falls  of  cast  Ranger  Canyon.  Mr.  L.  T.  Murray  espied  a  frog  in  amongst  the 
rocks  in  our  very  midst.  It  was  an  adult  female  Hyla  arenicolor. 

July  i  i,  Toyahvalc.  We  drove  to  Madera  Canyon  for  a  night  camp.  ...  In 
the  canyon  caught  a  recently  transformed  Hyla  aremcolor,  and  just  about  dark 
heard  several  males  calling. 

June  iH-2i,  11^4,  Pcna  Blanca  Spring,  Ariz.  Found  two  Hyla  aremcolor  in 
the  overflow  area  around  the  spring, 

April  26,  1942.  In  Santa  Anita  Canyon  (see  R.  b.  muscosa),  Calif.,  beside  the 
river  we  caught  two  Hyla  aremcolor.  The  first  one  I  spied  was  on  gravel  be- 
side stream  and  hopped  in.  It  was  very  light  in  color— much  granitic  rock  here, 
light  gray  in  color.  Almost  no  sediment  in  stream. 

April  $o.  At  mouth  Strawberry  Creek,  Calif.,  above  bridge  there  leaped  in 
from  a  boulder  a  female  H.  aremcolor. 

May  2.  North  of  Rmcon  at  about  2000  feet  in  a  little  stream  beside  the  road, 
shaded  by  live  oaks,  found  perched  on  a  boulder,  3  inches  from  each  other,  a 
male  and  female  H.  aremcolor.  They  were  in  full  sunlight. 

May  6,  Buckman  Spring,  Calif.,  to  bridge  3*4-4  miles  south.  During  the 
evening  rode  north  toward  Buckman  Springs,  hearing  several  isolated  calls 
which  took  me  some  time  to  locate.  Usually  they  would  quit  when  I  ap- 
proached. One  was  in  a  bush.  Finally  on  a  boulder  upside  of  stream  current 
saw  one  with  bubble  throat  inflated.  Hyla  arenicolor.  Must  have  been  ten 


294 


HANDBOOK  OF  FROGS  AND  TOADS 


males  scattered  along  the  creek.  Their  call  is  not  long,  not  so  grating  as  that 
of  H.  regilla,  neither  loud  nor  strident.  No  such  voluminous  call  as  Hyla 
versicolor. 

Authorities9  corner: 

G.  P.  Englehardt,  Ariz.,  1917,  pp.  5-6.        J.  R.  Slevin,  B.C.,  1928,  p.  112. 
S.  R.  Atsatt  in  T.  I.  Storer,  Calif.,  1925,     L.  T.  Murray,  Tex.,  1939,  p.  5. 

p.  209. 

Viosca's  Tree  Frog,  Whistling  Tree  Frog,  Whistling  Frog,  Bird-voiced  Hyla, 

Bird-voiced  Tree  Frog 

Hyla  avivoca  Viosca.  Plate  LXII;  Map  21. 

Range:  Southeast  Louisiana  (Florida  parishes  of  Louisiana)  to  Florida 
(O.  C.  Van  Hyning) ;  Georgia  (F.  Harper) ;  Henderson,  Tenn.  (Endsley) ; 
Kentucky  (Parker). 

Habitat:  Tupelo  swamps  in  the  valleys  of  rivers  and  smaller  streams  on  tu- 
pe^o  or  cypress  trees  and  on  buttonbush  (adapted  from  Viosca,  1928). 

"The  Apalachicola  drainage.  Known  from  Jackson  and  Liberty  counties. 
Habitat — Tupelo,  titi  and  cypress  swamps.  Not  rare,  but  infrequently  seen  be- 
cause of  the  nature  of  its  habitat"  (A.  F.  Carr,  Jr.,  Fla.,  1940^  p.  58) . 

Size:  Adults,  i%-i %  inches.  Males,  28-39  mm.  Females,  32-49  mm. 

General  appearance:  These  frogs  are  a  small  edition  of  our  common  tree 
toad,  Hyla  v.  versicolor.  They  may  be  brown,  green,  or  gray,  light  or  dark. 
The  arms  and  legs  are  distinctly  barred.  There  is  the  characteristic  light  yellow 
or  white  spot  below  the  eye.  The  color  in  the  groin  and  on  the  rear  of  the  legs 
is  a  pale  yellowish  green,  instead  of  orange  as  in  Hyla  v.  versicolor  males.  The 
throats  of  these  males  are  more  or  less  darkened  with  black  specks.  The  dark 
pattern  on  the  back  consists  of  a  bar  across  the  head  and  the  eyelids  and  an 
irregular  area  on  the  back,  the  bulk  of  the  pattern  being  near  the  rump.  The 
skin  is  moderately  smooth,  in  some  being  very  finely  granular.  In  appearance 
they  are  more  slender  than  H.  versicolor,  and  when  dark  brown,  as  we  first 
saw  them,  they  reminded  us  strongly  of  Hyla  fetnoralis. 

Color:  Male.  Top  of  head  rainette  green,  courge  green,  or  Scheele's  green, 
with  same  color  on  upper  eyelid  and  back  to  mid-back  spot,  and  on  either  side 
of  this  spot.  Interspaces  of  hind  legs  night  green,  courge  green,  or  rainette 
green.  The  mid-dorsal  spot  appears  dark  olive  or  deep  olive.  Under  the  lens  its 
background  is  black,  spotted  with  apple  green.  An  oblique  bar  of  similar  color 
extends  from  upper  eyelid  toward  meson  but  does  not  meet  its  fellow.  The 
color  just  above  the  vent  may  be  clear  yellow-green.  One  band  across  middle 
of  femur  is  chamois  or  vinaceous-buff.  A  line  of  mummy  brown  separates  dor- 
sal color  of  hind  leg  from  ventral  color.  Outer  half  of  rear  of  femur  and  rear 
half  of  dorsum  is  spotted  with  light  lumiere  green.  Bars  on  tibia  green  like 
back  of  body.  Bars  on  tarsus  and  foot  with  cream-buff.  Spot  below  eye  is  pale 


TREE  FROGS:  HYLIDAE 


295 


glass  green.  Back  of  eye  is  a  broad  black  vitta  extending  to  shoulder.  It  sur- 
rounds the  tympanum.  From  this  vitta  to  groin  is  a  row  of  black  spots  sepa- 
rating dorsum  from  venter.  Tympanum  is  more  or  less  the  same  color  as  femur 
bar  or  interspaces  on  top  of  arms  and  hands,  which  are  almost  old  gold.  Sub- 
orbital  spot  is  margined  with  mummy  brown  or  black.  The  forward  part  of 
chin  is  marked  with  black  dots  below  the  lower  labial  border.  Forward  throat 


•  +Hyla  avivoca9 1 


Map  21 

is  white  with  a  wash  of  marguerite  yellow.  Lower  throat  is  pale  flesh  to  Vene- 
tian pink.  Lower  belly  and  lower  surface  of  femur  are  the  same.  Rear  of  femur 
below  vent  and  rear  of  tarsus  are  marked  with  white  on  pebbly  surface.  Lower 
belly  and  groin  have  wash  of  viridme  green  or  other  yellow-green.  Underside 
of  fore  limb  is  like  fore  part  of  venter,  lichen  green  or  light  gull  gray.  Pupil  rim 
of  eye  citron  yellow  or  strontian  yellow;  ins  black,  heavily  spotted  with  these 
yellows. 

Female,  Mariana,  Fla.,  from  O.  C.  Van  Hyning,  Jan.  3,  1933.  Top  of  head 
drab,  buffy  brown  and  the  same  color  on  forward  part  of  dorsum.  The  bar 
from  eye  to  eye,  the  forward  dorsal  spot,  and  the  rear  dorsal  spot  olive-brown. 
The  side  of  body  has  background  pale  olive-buff  to  pale  vinaceous-fawn,  the 


296  HANDBOOK  OF  FROGS  AND  TOADS 

groin  olive-buff  or  pale  glass  green.  The  light  interspace  color  on  rear  of  femur 
may  be  glass  green,  pale  glass  green,  or  pale  olivine.  The  transverse  bars  on 
tibia,  interspace  color  on  rear  of  femur,  and  spots  on  groin  are  bister  or  raw 
umber.  The  pale  spot  below  the  eye  is  clear,  prominent,  pale  olive-buff  or 
marguerite  yellow.  The  whole  chin  is  white  with  very  few  black  dots  near 
lower  labial  border.  The  under  parts  in  general  are  white  or  pale  olive-buff, 
with  scarcely  any  suggestion  of  the  wash  of  greenish  present  in  the  male.  The 
pupil  rim  is  encircled  by  a  carnelian  red  or  vinaceous-tawny  ring;  the  iris 
proper  is  black  dotted  with  testaceous  or  terra  cotta  spots. 

Structure:  More  slender  than  Hyla  versicolor;  muzzle  truncate;  webs  large, 
but  leaving  the  last  two  joints  of  fourth  toe  free,  except  for  web  margin;  back 
almost  smooth;  ventral  surface  with  distinct  granulations;  usually  with  nar- 
row waist;  male  with  folds  on  rear  of  throat  almost  to  pectoral  region. 

Voice:  "The  voice  is  birdlike,  being  a  plaintive  whistle  repeated  in  quick 
succession,  much  as  in  the  redbellied  woodpecker.  This  call  is  sometimes  pre- 
ceded by  a  few  notes  of  a  slower  call  much  like  the  voice  of  Hyla  crucifer. 

"The  pitch  and  tone  of  the  voice  of  avivoca  is  nearer  to  that  of  crucifer  than 
to  any  other  eastern  American  Hyla  although  its  rate  is  far  more  rapid  than 
that  of  crucifer"  (P.  Viosca,  Jr.,  La.,  1928,  pp.  90-91). 

"The  males  call  from  bushes  in  or  near  the  water  and  from  cypress,  tupelo, 
and  black-gum  boles"  (A.  F.  Carr,  Jr.,  Fla.,  1940^  p.  58). 

See  F.  Harper,  Gen.,  1935,  pp.  290-293,  for  characteristics  of  its  call. 

Breeding:  This  species  breeds  from  June  to  mid-August.  The  eggs  and  tad- 
poles are  not  described.  "Hyla  avivoca  is  more  clearly  related  to  Hyla  versicolor 
than  to  any  other  North  American  Hyla.  .  .  ." 

"I  located  a  small  chorus  in  Sweetwater  Creek  Swamp,  Liberty  County, 
April  19,  1935,  where  H.  crucifer,  H.  v.  versicolor,  Pseudacns  jenartim,  and 
Rana  clamitans  were  also  calling.  I  have  never  seen  a  female"  (A.  F.  Carr,  Jr., 
Fla.,  i94ob,p.  58). 

Journal  notes:  June  8, 1930,  New  Orleans,  Sunday.  Went  to  Mandevillc  and 
Lewisburg  by  myself  to  search  out  Viosca's  type  locality.  Didn't  succeed. 

June  9,  New  Orleans.  Went  to  Viosca's  house.  Heard  H.  c.  ctnerea.  While 
there  Chase  from  office  called  up  that  by  dint  of  hard  work  he  had  taken  four 
Hyla  avivoca  at  Lewisburg,  Covington,  and  in  a  swamp  just  south  of  Mande- 
ville.  They  are  males.  They  surely  are  a  different  frog. 

June  10,  New  Orleans.  Started  at  noon  for  the  north  country,  Pearl  River, 
and  pine  barren  parishes  of  Louisiana,  with  Percy  Viosca,  Jr.,  and  H.  B.  Chase. 
Went  to  Pearl  River.  Didn't  hear  H.  avivoca  in  the  gum  and  cypress  edges,  a 
haunt  of  the  species.  Presently  we  heard  a  Flycatcher,  which  I  remarked  was 
like  a  crested  Flycatcher.  Then  Chase  got  excited  over  a  note  in  the  tupelo 
swamp  either  side  of  the  bridge.  Instantly  the  setting  and  note  made  me  say, 
"It  is  a  Pileated  Woodpecker."  A  day  or  so  before  Viosca  and  Chase  had  said 


TREE  FROGS:  HYLIDAE 


297 


Plate  LXII.  Hyla  avivoca  (y(iVi).  1-4. 
Males. 


Plate  LX1II.  Hyla  baudinii  (X%).  i.  Fe- 
male. 2-7.  Males. 


298  HANDBOOK  OF  FROGS  AND  TOADS 

this  frog's  note  was  like  a  Pileated  Woodpecker  at  a  distance.  Naturally  they 
both  laughed  at  me.  Worth  it.  Would  I  have  guessed  it  to  be  a  frog,  much  less 
a  Hyla?  Fear  not.  Heard  a  few  of  them  in  the  gums  of  Tick  Faw  River  where 
we  will  go  tomorrow  night. 

June  ii.  Tick  Faw  River,  La.  (near  Ferry).  In  Tick  Faw  River  we  heard  a 
few  H.  avivoca.  Finally  Chase  called  me  to  hurry  and  started  off  on  the  run. 
I  followed.  We  were  soon  in  a  tupelo  swamp.  Near  the  edge  we  heard  them 
Chase  found  his  first  one  crosswise  of  leaves  of  buttonbush  (Cefhalanthus 
occidentalis) .  My  first  was  on  the  bole  of  a  gum.  Many  of  Chase's  captures 
were  head  down  on  upright  branch  of  buttonbush.  Mine  were  head  up.  Chase 
and  I  followed  voices.  We  got  five  frogs  between  us.  Viosca  with  a  lamp  on 
his  head  shone  their  eyes.  He  didn't  follow  voices  so  much.  He  got  more  in 
this  way  than  either  of  us.  One  frog  was  pure  green  with  no  markings.  Viosca 
found  several  on  small  gums  near  the  tupelo  edge  while  Chase's  and  my  cap- 
tures were  farther  in  the  swamp.  On  the  trees  was  some  poison  ivy.  On  clumps 
or  tree  bases  in  the  swamp  and  at  the  edge  of  the  swamp  were  sensitive  fern 
(Onoclea  senstbilis),  royal  fern  (Qsmunda  regalis),  Utrictdaria  (purple),  a 
marsh  St.-John's-wort,  sphagnum,  water  penny  (Hydrocotyle),  lizard's-tail 
(Saururus),  Nyssa  aquatica,  and  buttonbushes  with  mayhaws.  Most  of  the 
frogs  were  3  or  4  feet  above  the  water.  Finally  Chase  found  a  female  on  a  log, 
in  the  water.  Sometimes  they  go  higher  in  trees.  Usually  they  are  down  at  3-4 
feet  or  less.  Their  call  is  birdlike.  It  is  a  fine,  delicate  little  species.  Viosca  did 
right  to  call  it  a  new  form,  H.  avivoca. 

June  ii.  Tick  Faw  River,  La.  Throat  in  croaking  is  somewhat  swollen,  vi- 
brates for  6-10  or  12  calls.  These  are  more  or  less  the  same,  repeated.  Sometimes 
raises  rear  end  with  calling.  Chase  also  noticed  the  vibration  or  varied  swelling 
at  each  call. 

Authorities9  corner: 
P.  Viosca,  Jr.,  La.,  1928,  p.  91. 
J.  R.  Endslcy,  Tenn.,  1937,  p.  70. 

Mexican  Tree  Frog,  Van  Vliet's  Frog 

Hyla  battdinii  (Dumcril  and  Bibron) .  Plate  LXIII;  Map  20. 

Range:  South-central  and  southern  Texas  through  Central  America. 

"This  Mexican  species  has  been  found  by  Mr.  Marnock  in  the  low  country 
southwest  of  San  Antonio,  commencing  with  San  Miguel  Creek,  a  tributary 
of  the  Medina.  This  is  its  most  eastern  known  range,  that  previously  given  by 
Professor  Baird  being  Brownsville  (as  Hyla  vanvlietif)"  (E.  D.  Cope,  Tex., 
i88oa,  p.  29). 

Habitat:  June  18, 1930.  At  10  P.M.,  two  miles  west  of  Brownsville,  in  a  resaca, 
found  these  frogs  in  small  bushes,  in  weedy  clumps,  and  even  in  grassy  tangles 
in  overflowed  tomato  field  adjoining  the  overflowed  resaca.  Later,  along  the 


TREE  FROGS:  HYLIDAE 


299 


Rio  Grande  in  Brownsville,  found  them  in  a  pond.  Mr.  Blanchard  tells  me  he 
saw  Mr.  Camp  take  them  along  the  river  (Rio  Grande)  in  palms  just  above 
Mr.  Rebb's  palm  grove. 

June  19.  Heard  several  in  the  palm  grove  which  is  completely  flooded  by 
high  river.  Beyond  grove,  and  along  river,  a  large  chorus  is  calling. 

Size:  Adults,  1/4-3%  inches.  Males,  44-71  mm.  Females,  44-89  mm. 

General  appearance:  This  large  tree  frog  has  a  black  patch  over  the  arm 
insertion  and  a  white  line  encircling  the  arm  insertion.  Its  color  ranges  from 
nearly  black  to  light  yellow  green,  gray,  or  fawn.  It  has  a  transverse  bar  be- 
tween the  eyes.  Various  irregular  lateral  spots  of  black  form  a  reticulation  on 
a  yellow  or  olivine  side  area.  There  are  transverse  bars  on  the  legs,  a  light 
greenish  spot  under  the  eye,  a  light  line  above  the  dark-edged  upper  jaw,  and 
a  dark  line  from  eye  to  shoulder  ending  in  a  prominent  black  patch.  The 
breast  and  throat  may  be  spotted  with  dark.  The  sides  are  yellow  posteriorly. 
In  the  female,  the  under  parts  of  the  throat  and  upper  breast  are  white,  the 
lower  breast  and  belly  white  suffused  with  pale  green.  In  the  male,  under 
parts  arc  cream  buff  with  lateral  throat  sacs,  a  pale  brownish  drab.  The  rear  of 
the  femur  is  yellow  and  russet. 

Color:  Male.  Brownsville,  Tex.,  June  19,  1930.  Back  sulphine  yellow  and 
aniline  yellow  on  side  or  olive  lake  on  back  becoming  old  gold  on  sides  (no 
dorsal  marks;  last  night  this  frog  was  brown  with  prominent  marks  on  dor- 
sum).  The  back  may  be  chamois  to  primrose  yellow  with  olive-yellow  or 
some  green  interspersed.  Cream  buff  line  along  jaw  to  beyond  angle  of  mouth 
and  under  tympanum.  Tympanum  pinkish  cinnamon  or  light  vinaccous- 
cinnamon  or  vinaceous-buff.  From  eye  over  tympanum  irregular  black  line 
ending  in  a  large  vertical  curl  of  black  above  arm  insertion.  Along  side  and 
a  little  below  level  of  black  arm  insertion  spot  are  several  prominent  black 
spots  to  the  groin.  This  black  spotting  is  more  or  less  reticulated  with  lemon 
yellow.  Top  of  fore  limbs  warm  buff,  pinkish  buff,  or  chamois.  From  eye  al- 
most to  black  shoulder  spot  is  a  wood  brown  band.  Chin  sulphur  yellow  or 
light  green-yellow.  Each  lateral  sac  with  oil  yellow  to  ecru-drab  or  pale  brown- 
ish drab.  Under  parts  cream  buff  except  a  narrow  pectoral  area  and  extension 
between  two  sacs.  This  little  area  is  white.  Hind  legs  more  or  less  color  of 
forelegs.  Rear  of  femur  with  some  viridine  yellow  or  bright  green-yellow  with 
vinaceous-russet;  under  side  of  hind  legs  onion-skin  pink  with  every  tubercle 
crowned  with  sulphur  yellow.  All  of  venter  very  pebbly.  Iris  very  bronzy; 
below  heavily  spotted  with  buff-pink,  above  it  is  chartreuse  yellow;  pupil 
rim  pale  greenish  yellow,  broken  in  front  and  behind;  outside  iris  is  ring  of 
lichen  green  or  Niagara  green. 

Other  males.  One  greenish  yellow  on  back  except  for  top  of  head  and  top 
of  hind  legs  and  forelegs,  which  are  viridine  yellow.  Another  bright  green- 
yellow  on  back.  From  eye  of  each  extends  a  horizontal  black  line  ending  in  a 
vertical  black  curve  back  of  axil.  Below  this  is  a  clay  color  or  cinnamon  band 


300  HANDBOOK  OF  FROGS  AND  TOADS 

outlined  by  black  above,  and  below  by  light  line  from  upper  jaw  edge  almost 
to  base  of  vertical  black  bar.  Tympanum  may  be  vinaceous-cinnamon  or  cin- 
namon. Along  side  where  green  of  back  ends  and  before  greenish  yellow  of 
reticulated  or  spotted  side  begins  is  cinnamon-buff.  In  one,  no  black  spots  and 
little  yellow.  Iris,  upper  part  cinnamon-buff. 

Female.  Brownsville,  Tex.,  from  H.  C.  Blanchard,  July  23,  1930.  The  for- 
ward part  of  head,  upper  eyelid,  face,  and  arms  are  vinaceous-buff.  The  back- 
ground of  back  and  legs  is  deep  olive-buff  or  olive-buff.  The  bars  of  tibia, 
femur,  and  arm,  the  dark  spots  on  back,  and  the  large  spot  from  each  eyelid 
on  fore  back  are  grayish  olive  or  vetiver  green.  Particularly  in  the  groin  and 
slightly  on  sides  are  black  spots  on  pale  olivine  ground.  The  reticulation  of 
the  sides  is  less  prominent  than  in  the  male,  and  restricted  almost  entirely  to 
the  groin.  The  black  U-shaped  spot  on  shoulder  has  the  opening  toward  the 
tympanum.  The  tympanum  is  wood  brown.  The  iris  is  somewhat  duller  than 
in  the  male;  lower  ins  wood  brown,  the  upper  avellaneous.  The  under  parts 
of  hands  are  light  vinaccous-fawn.  The  under  parts  of  feet,  femur,  and  tibia 
are  light  russet-vinaceous.  The  under  parts  of  throat  and  upper  breast  are 
white;  the  lower  breast  and  belly  are  white  with  a  suffusion  of  pale  turtle 
green.  After  the  frog  had  been  in  a  moist  jar  for  a  few  hours,  the  back  became 
darker,  Prout's  brown,  and  the  reticulation  along  the  sides  became  more 
marked. 

Aug.  13,  the  female  still  continues  with  the  rich  brown  coloration.  She  re- 
mains under  the  wet  moss.  The  two  males  come  up  in  the  glass  jar,  one  is  a 
bright  green  and  the  other  old  gold  or  citrine.  Female  54  mm. 

Structure:  Skin,  smooth  above,  set  with  fine  tubercles;  under  parts  granu- 
lar; tympanum  nearly  as  large  as  eye;  fold  of  skin  from  eye  to  shoulder; 
prominent  fold  across  breast;  disks  large,  fingers  slightly  webbed,  toes 
webbed;  a  distinct  tarsal  fold. 

"Hyla  vanvlieth  Baird — Nearly  smooth  above.  Tympanum  nearly  as  large 
as  the  eye.  Tibia  half  as  long  as  the  body,  longer  than  arm  from  elbow,  which 
in  turn  exceeds  the  foot.  Ash  gray  or  olive,  with  an  irregular  cruciform  dorsal 
blotch.  Thigh  and  leg  with  three  transverse  bands  each.  Their  inner  surfaces 
when  flexed  scarcely  reticulated,  but  spotted  with  white  upon  a  darker 
ground.  Inside  of  tibia  uncolored.  Body  two  inches.  Hab.  Brownsville,  Texas" 
(S.  F.  Baird,  Gen.,  1856,  p.  61). 

Voice:  The  vocal  sac  is  better  developed  on  either  side  of  throat  than  in  its 
middle.  This  is  especially  noticeable  in  the  live  frog. 

June  18, 1930,  Brownsville,  Tex.  The  first  time  I  heard  them  I  guessed  they 
must  be  H.  baudinti.  Heard  their  chorus  at  %  mile.  The  note  was  a  blurred 
Aec^  or  l(ecl(  (not  high  like  Acris).  This  may  be  repeated  5  to  8  or  even  10  or 
12  times.  Then  comes  an  interesting  chuckle,  or  no  chuckle  at  all,  or  no  fecl^ 
and  all  chuckle.  Never  heard  anything  like  it.  The  first  call  (not  chuckle)  in 
some  ways  reminds  me  of  Hyla  cinerea,  but  yet  it  is  much  unlike  the  call  of 


TREE  FROGS:  HYLIDAE  301 

the  cowbell  tree  frog.  Does  the  full  dilation  of  the  sac  on  either  side  cause  the 
queer  note  (unlike  most  Hylas)  ? 

In  Mr.  F.  Rebb's  palm  grove  heard  a  few  H.  batidinu.  They  are  too  far  out 
in  an  overflow  to  reach  them.  To  the  east  of  his  house  and  near  the  river  heard 
a  chorus  of  these  frogs.  Their  repeated  fecfo  or  hect(s  last  about  2  or  3  seconds, 
then  as  much  interval,  whereas  B.  vdliceps  has  a  longer  call,  sometimes  to  5 
seconds. 

Breeding:  The  only  record  on  transformation  is  of  one  specimen  trans- 
formed, or  just  past,  from  Panama,  %  inch  (21  mm.)  in  length,  which  was 
caught  Feb.  19,  1911.  They  were  in  full  chorus  in  Brownsville,  Tex.,  June  19, 
1930.  In  southern  Vcra  Cruz,  A.  G.  Ruthven  found  this  species  common.  He 
observed  them  breeding  on  July  17.  In  1908,  near  Cordoba,  Vera  Cruz,  Dr.  H. 
Gadow  found  a  spawning  congress  of  45,000  frogs.  His  account  is  very  vivid. 

"Whilst  rambling  along  the  edge  of  the  forest  we  became  conscious  of  a 
noise,  at  first  resembling  the  mutter  of  a  distant  sawmill;  but  on  our  reaching 
the  other  side  of  a  cluster  of  trees  this  sound  grew  into  a  roar,  like  that  of 
steam  escaping  from  many  engines,  mingled  with  the  sharp  and  piercing 
scream  of  saws.  It  came  from  a  meadow  containing  a  shallow  pool  of  rain- 
water. In  the  wet  grass,  on  its  stalks,  and  on  the  ground,  hopped  about  hun- 
dreds of  large  green  tree-frogs;  nearer  the  pool  they  were  to  be  seen  in  thou- 
sands, and  in  the  water  itself  there  were  tens  of  thousands.  Hopping,  jumping, 
crawling,  sliding,  getting  hold  of  each  other,  or  sitting  still.  Most  of  them 
were  in  amplexits,  and  these  couples  were  quiet,  but  the  solitary  males  sat  on 
their  haunches  and  barked  solemnly,  with  their  resounding  vocal  bags  pro- 
truding. Every  now  and  then  one  was  making  for  a  mate,  and  often  there 
were  three  or  four  hanging  on  to  each  other  and  rolling  over.  The  din  was  so 
great  that  it  was  with  difficulty  that  we  caught  the  remarks  that  we  shouted, 
although  we  were  standing  only  a  iew  feet  apart.  Each  sweep  of  a  butterfly 
net  caught  at  least  half-a-dcr/cn  frogs. 

"Now  the  grassy  pool,  where  the  frogs  were  closest,  was  about  30  yards 
square  (900  square  yards),  rather  more  than  the  area  of  a  tennis  lawn,  and 
each  square  yard  held  from  50  to  100  frogs — many  square  yards  certainly  held 
several  hundreds  each.  At  the  very  lowest  computation  this  gives  45,000  frogs; 
and  there  was,  besides,  an  outer  ring  of  some  five  hundred  square  yards  where 
frogs  were  fairly  numerous,  say  from  5  to  10  to  the  square  yard,  mostly  spent 
females,  but  these  few  thousands  we  may  leave  out  of  the  reckoning,  to  under- 
state rather  than  overestimate  the  number.  Supposing  there  were  only  20,000 
females,  each  spawning  from  5,000  to  10,000  eggs — say  only  5,000 — the  total 
would  amount  to  just  100,000,000  eggs.  The  spawn  literally  covered  both 
ground  and  water  thickly.  But  the  greatest  surprise  awaited  us  on  the  fol- 
lowing morning,  when  we  went  to  photograph  the  scene.  There  was  not  a 
single  frog  left;  the  water  had  all  evaporated,  and  the  whole  place  was  glazed 
over  with  dried-up  spawn!  The  prospective  chance  of  millions  of  little  frogs 


302  HANDBOOK  OF  FROGS  AND  TOADS 

was  gone,  their  expectant  parents  having  been  deceived  in  calculating  their 
day  of  incarnation.  That  was  on  the  4th  of  July,  several  weeks  after  the  be- 
ginning of  the  rather  fitful  rainy  season"  (H.  Gadow,  Gen.,  1908,  pp.  75-76). 

"We  found  H.  bandinii  common  at  Cuatotolapam.  Most  of  the  specimens 
were  taken  during  night  rains  on  the  banana  trees  at  San  Juan.  At  these  times 
they  were  very  noisy.  During  the  clay  we  found  them  secreted  under  boards, 
in  the  bases  of  such  large  leaved  plants  as  the  'elephant  ears,'  bananas,  etc. 
They  were  observed  breeding  in  a  pond  near  La  Laja  Creek  on  July  17" 
(A.  G.  Ruthven,  Gen.,  1912,  pp.  310-311). 

On  the  basis  ot  some  material  from  Mr.  Sumichrast  (taken  at  Tehuantepec, 
Mexico)  Cope  (Gen.,  i88ob,  p.  267)  wrote:  "Abundant,  but  only  seen  in  the 
rainy  season,  when  it  comes  to  pools,  lagoons,  etc.  to  breed." 

On  Dec.  31,  1952,  Prof.  H.  J.  Swanson  of  Edinburg,  Tex.,  sent  us  three 
specimens,  30,  32,  and  32  mm.,  respectively.  They  had  the  reticulated  sides, 
dark  spot  below  eye,  bar  on  shoulder  insertion,  and  white  stripe  across  vent 
with  black  spot  below. 

Journal  notes:  June  18,  1930,  Brownsville,  Tex.  That  night,  went  out  the 
Military  Road  2  or  3  miles  where  the  road  comes  opposite  a  rcsaca  which  is 
%-%  mile  from  the  road.  From  road  heard  a  chorus  new  to  me.  My  guess 
was  H.  baudinii.  Pell-mell  I  started.  The  resaca  looked  very  foul  and  dirty 
because  of  ripe,  overripe,  and  green  tomatoes  afloat.  On  little  bushes  or  among 
tomato  vines  heard  these  new  frogs.  The  note  was  a  heavy  f(ec^  given  5-10 
times  with  an  interesting  chuckle  following.  The  first  frog  heard  I  never 
could  locate,  though  I  was  almost  on  it,  in  such  a  tangle  of  grass  and  vines. 
Finally  located  one  in  a  bush  where  water  was  waist-deep.  It  was  on  a  branch 
i  foot  above  water.  This  one  was  yellow-green.  Then  followed  another.  It 
was  brown  in  color  and  on  the  ground  where  the  water  was  shallow.  Tried 
for  several  more.  Whenever  I  approached,  they  stopped.  Did  my  white  shirt 
or  something  cl&c  give  me  away?  For  some  time  raced  back  and  forth  be- 
tween two  Hyla  buudtmi  and  three  narrow-mouthed  toads.  Didn't  get  the 
two  Baudm's  frogs.  After  i%  hours'  effort  turned  back  for  fear  taximan  had 
gone  or  thought  me  lost.  By  pointers  (stars)  worked  my  way  back.  Came  out 
by  the  car.  In  a  pond  beside  the  river  heard  another  H.  bandmri.  It  was  deep 
green  on  back.  Only  the  brown  specimen  had  markings  on  back.  The  yel- 
lowish green  and  deep  green  specimens  were  all  green  on  the  back.  The  last 
one  was  found  on  the  surface  of  the  water  among  a  clump  of  weeds. 

August  22,  1930.  Went  over  to  Mrs.  Olive  Wiley's  museum  in  the  Min- 
neapolis Public  Library.  She  had  a  beautiful  Hyla  baudinii  from  Central 
America.  It  came  on  a  bunch  of  bananas  to  a  local  merchant. 

Authorities'  corner: 

A.  Gunther,  Gen.,  1885-1902,  pp.  271-272.          R.  Kellogg,  Gen.,  1932,  p.  161. 
J.  K.  Strecker,  Jr.,  Tex.,  1915,  p.  50.  W.  G.  Lynn,  Gen.,  1944,  p.  189. 


TREE  FROGS:  HYLIDAE 


303 


Green  Tree  Frog,  Carolina  Tree  Frog,  Marsh  Tree  Frog,  Cinereous  Frog, 
Bell  Frog,  Fried  Bacon  Frog,  Cowbell  Frog,  Bull  Frog 

Hyla  cinerea  cinerea  (Schneider).  Plate  LXIV;  Map  22. 

Range:  Virginia  to  Texas,  up  the  Mississippi  River  to  Illinois  and  Missouri. 

Habitat:  Swampy  edges  of  watercourses;  on  the  taller  water  plants  in 
ditches  or  pools;  on  lily  pads,  trees,  bushes,  or  vines  not  far  from  water. 
Prof.  E.  A.  Andrews  in  1928  reported  finding  them  in  pitcher-plant  trumpets. 


Map  22 

The  frogs  may  be  on  the  bushes  and  stems  above  the  water,  but  more  fre- 
quently are  at  the  water's  level.  During  the  day  they  rest  mainly  on  the  stems 
above  the  water.  They  may  be  hidden  in  clumps  of  Spanish  moss  or  under 
flakes  of  bark  on  trees.  June  27, 1922,  at  Camp  Pmckncy,  we  took  Hyla  cinerea 
in  Pond  No.  2,  on  a  vertical  wild  rice  stem,  the  frog  some  3  feet  above  the 
water.  July  5,  at  Pond  No.  3,  on  the  wide-leaved  grasshkc  plants  were  some 
7  or  8  Hyla  cinerea  perched  1-3  feet  above  the  water.  On  August  u,  found 
3  males,  2  feet  above  the  water. 

"The  green  tree  frog  has  been  found  climbing  out  on  the  emergent  portions 
of  water  hyacinths  on  several  occasions.  Concerning  this  species,  Kilby  (MS 
1938)  says:  'By  far  the  greatest  concentration  of  individuals  is  found  among 


304  HANDBOOK  OF  FROGS  AND  TOADS 

lake  and  pond  shore  vegetation  and  on  the  floating  rafts  of  water  hyacinths 
that  are  common  in  Florida  Lakes.'  My  own  records  are  for  January,  Febru- 
ary, and  October"  (C.  J.  Coin,  Fla.,  1943,  pp.  148-149). 

"This  is  the  most  beautiful  tree-frog  of  our  fauna.  It  lives  on  the  leaves  of 
plants,  frequenting  especially  lily  pads  and  other  aquatic  vegetation  at  the 
edges  of  lakes.  It  occurs  also,  at  times,  in  fields  of  corn.  Its  food  consists  of 
insects,  the  common  fly  being,  it  is  said,  preferred.  Its  note  resembles  the  tone 
of  a  cow  bell  heard  at  a  distance.  Where  abundant  about  water,  the  frogs  are 
very  noisy  just  before  dusk,  the  chorus  being  broken,  however,  by  longer  or 
shorter  intervals  of  silence.  A  single  note  is  first  heard,  and  as  if  that  were  a 
signal,  it  is  taken  up  and  repeated  by  a  dozen  noisy  throats  till  the  air  is  reso- 
nant with  the  sound.  After  a  time  it  ceases  as  suddenly  as  it  began,  to  be  again 
resumed  after  a  period  of  quiet"  (H.  Garman,  111.,  1892,  pp.  347-348). 

Size:  Adults,  i^-a1/^  inches.  Males,  37-59  mm.  Females,  41-63  mm. 

General  appearance:  This  is  a  very  slim,  smooth,  bright  green  tree  frog  with 
a  light  side  stripe,  pointed  head,  and  shallow  face.  It  is  relatively  the  longest- 
legged  Hyla  of  the  East.  Often  there  are  small  gold  or  yellow  spots  on  the 
back.  The  under  surfaces  are  white  or  yellowish  white. 

Deckert  (Fla.,  1915,  p.  3)  described  it  as  "an  aristocratic  looking  tree  toad, 
with  its  long,  slender  figure  of  the  brightest  green,  edged  on  each  side  with 
a  band  of  pale  gold  or  silvery  white." 

"This  is  a  larger  species  than  Hyla  lateralis,  the  length  of  which  according 
to  Daudin,  is  *un  pouce  et  demi  au  plus.'  Dr.  Holbrook's  specimen,  however, 
measured  i%  inches.  The  largest  specimen  in  the  collection  of  the  Academy 
measures  i|X>  inches  (Fr.).  It  is  a  much  more  slender  animal  than  semifas- 
ciata.  In  laterahs  (viridis,  Holl.)  the  lateral  stripe  extends  as  far  as  the  anus, 
and  there  is  a  white  band  running  the  whole  length  of  the  tibia,  both  ante- 
riorly and  posteriorly.  The  anterior  band  is  absent  in  semifasciata"  (E.  Hallo- 
well,  Tex.,  i857b,  pp.  307-308). 

Color:  Okefinokee  Swamp,  Ga.,  April  26,  1921.  Male.  In  dark  olive-green 
males  there  are  twenty  orange  spots  scattered  over  the  back.  The  frog  is 
lighter  green  on  sides  below  lateral  stripe,  which  is  cream-colored  and  ex- 
tends almost  to  hind  legs.  Obscure  on  snout  ahead  of  eye.  Cream  colored  or 
white  stripe  on  back  of  forearm.  Same  on  back  of  lower  leg  and  along  hind 
end  of  foot.  Throat  from  angle  of  mouth  to  slightly  back  of  chin  green.  Chin 
proper  yellowish  cream.  Back  of  these  two  areas,  yellowish  cream  and  green, 
is  the  wrinkled  pink  area  of  the  throat  proper.  One  light-colored  male  has 
stripe  straw-colored  and  extending  only  an  inch  behind  insertion  of  arm  area. 
There  is  black  above  and  below  the  stripe.  (Non-Ridgway.) 

Another  male.  Back  may  be  apple  green,  dark  olive-green,  greenish  olive,  or 
deep  slaty  olive  to  almost  black,  the  stripe  on  side  light  dull  green-yellow  or 
clear  dull  yellow.  In  one  male,  stripe  not  beyond  tympanum  and  suggestive 


TREE  FROGS:  HYLIDAE 


305 


of  H.  evittata.  The  vent,  forearm  stripe,  foot  stripe,  and  heel  are  white.  One 
dark  male  had  the  foot  stripe  fainter.  Iris  russet-vinaceous. 

Female.  Has  no  pink  chin,  and  green  extends  in  on  sides  of  throat  for  slight 
distance.  Straw-colored  stripe  to  insertion  of  hind  legs  prominent,  also  stripe 
under  eye.  Under  parts  from  groin  to  chin  the  same  cream  color.  (Non- 
Ridgway.) 

Structure:  Slender,  flat  in  body;  skin  smooth  or  minutely  granular;  breast 
fold  present;  vocal  sac  a  round  subgulai  pouch;  a  tympanic  fold  from  tympa- 
num to  base  of  arm. 

Voice:  The  voice  is  loud  and  at  a  distance  sounds  like  a  cowbell.  The  indi- 
vidual call  is  quon/t,  qnonl{,  quvn\,  qnan^  To  some  ears,  fried  bacon,  fried 
bacon.  To  Deckert,  grab,  grab,  grabit,  grabit.  This  is  easily  one  of  the  most 
characteristic  anuran  voices  of  the  South.  It  is  one  of  the  rain  signals  for  the 
residents,  who  call  it  the  "ram  frog."  In  the  daytime  when  the  weather  is 
sultry  or  especially  in  the  evenings  of  late  May  or  early  June,  some  of  the 
immense  choruses  are  not  easily  to  be  forgotten.  Sometimes  a  chorus  starts 
suddenly,  quickly  reaches  its  crest,  and  ends  abruptly,  to  be  resumed  later 
after  a  shorter  or  longer  sharply  defined  interval.  Along  some  of  the  water- 
courses like  Billy's  Lake  one  lone  frog  near  by  will  begin,  then  stop,  but  before 
he  has  finished,  another  just  ahead  of  the  speeding  boat  has  taken  up  the  task. 
Thus  the  chorus  may  travel  along  the  margin  of  a  lake  for  considerable  dis- 
tances. 

At  Flatwood,  Ala.,  1917,  after  6:30  P.M.  in  a  drying-up,  swampy  pond,  we 
heard  a  chorus  which  sounded  to  one  member  of  the  party  like  an  exhaust 
running  into  an  oil-well  pipe.  We  took  one  frog  with  the  aid  of  a  flashlight. 
It  was  on  some  bushes.  Its  sides  and  throat  looked  like  a  pink  ball. 

The  calls  may  be  given  about  75  to  the  minute.  The  frog  may  redouble  its 
speed  and  at  the  same  time  add  a  rolling  quality  to  the  note:  crron\ — crron^ 
— crronf{.  Two  individuals  calling  together  but  not  in  unison  produce  bo  babe, 
bo  babe,  bo,  babe. 

J.  Le  Conte  (Gen.,  1856,  p.  428)  said  it  was  commonly  known  as  the  bell 
frog,  its  notes  resembling  the  sound  of  small  bells. 

J.  E.  Holbrook  (Gen.,  1842,  IV,  120-121)  wrote:  "Their  noise  proceeds  from 
a  single  note,  which  at  a  little  distance  is  not  unlike  the  sound  of  a  small  bell; 
and  there  seems  in  general  to  be  one  leader  of  their  orchestra,  and  when  he 
raises  his  note,  hundreds  take  it  up  from  all  parts  of  the  corn  field,  and  when 
he  stops,  the  concert  is  at  an  end,  until  he  again  begins." 

Breeding:  This  species  breeds  from  April  15  to  August  15.  The  black  or 
brownish  and  white  or  cream  eggs  are  in  small  packets  or  films  at  or  near  the 
surface,  attached  to  floating  vegetation.  The  outer  envelope  is  poorly  defined, 
becoming  part  of  the  mass.  The  egg  is  Ho-Vio  inch  (0.8-1.6  mm.),  the  inner 
envelope  V\^r%  inch  (2.2-3.4  mm.),  the  outer  envelope  %-%  inch  (3.6-4.0 


306  HANDBOOK  OF  FROGS  AND  TOADS 

mm.).  The  tadpole  is  medium,  i%  inches  (40  mm.),  its  tail  long  and  acumi- 
nate, its  body  green  with  a  sulphur  or  ivory  stripe  on  the  side  of  the  head  from 
snout  to  eye.  The  tooth  ridges  are  %.  After  55  to  63  days,  the  tadpoles  trans- 
form from  July  2  to  October,  at  V&-1 1A  o  inch  ( 12-17  mm.). 

"The  breeding  season  extends  from  late  spring  to  late  summer.  .  .  .  The 
condition  of  the  weather  is  a  controlling  factor  in  determining  when,  within 
this  period,  breeding  actually  occurs"  (J.  D.  Kilby,  Fla.,  1945,  p.  82). 

Journal  notes:  The  same  conditions  do  not  always  produce  the  same  colora- 
tion. In  one  case,  we  had  many  Hyla  ctncrca  m  a  botany  drum.  All  were  light 
green  except  three,  one  of  which  was  almost  black,  another  olive  green  or  dark 
green,  and  the  third  yellowish  green.  One  noon  in  1917  we  stopped  in  a  low 
woods  near  one  of  the  bays  between  Pass  Christian  and  Bay  St.  Louis,  Miss. 
On  the  saw  palmetto  leaves  we  found  no  end  of  Hyla  cmerea  and  Hyla  squi- 
rella  in  all  color  phases.  These  creatures  were  on  the  leaves  or  in  the  bases 
between  two  stems.  In  one  place  found  two  specimens,  one  a  brown  phase 
and  one  a  green  phase.  Some  have  a  yellowish  or  orange  color  on  the  posterior 
faces  of  thighs,  others  have  purplish.  Some  have  yellowish  line  on  lip  and  a 
few  beside*  have  faint  yellowish  line  on  side.  May  have  a  purplish  area  from 
nostril  to  femur  along  the  side.  In  one  clump  one  yellowish  green  on  back, 
another  dark  greenish,  and  a  third  purplish  brown.  There  were  unspotted 
//.  evittiita;  Hyla  cinerea,  olive,  green,  or  brownish  with  or  without  lateral 
bands,  with  or  without  dark  borders  to  band. 

May  io,  1921.  This  evening  at  X  P.M.  (temperature  72°)  went  to  new  pond. 
.  .  .  Heard  a  Hyla  cinerea  near  by.  Hcgan  to  search.  It  stopped.  Later  thought 
I  saw  it.  Sci/cd  it  and  it  proved  a  ripe  female,  not  croaker.  This  female  was 
in  a  cypress  4  feet  above  the  water.  Went  away.  Later  the  croaker  started  again. 
On  the  flat  side  of  an  ins  leal  $  feet  from  the  captured  female  was  the  male. 

At  night  many  individuals  arc  seen  to  be  fairly  covered  with  certain  tiny 
insects  that  arc  common  in  the  ground  vegetation.  Some  that  were  collected 
proved  to  be  harmless  flics  (Osama  longgpes).  Their  perching  on  the  frogs  is 
probably  accidental. 

June  14,  im»f  Bccville,  Tex.  All  of  a  sudden  several  Hyla  cinerea  began 
croaking  around  the  pond.  They  were  on  near-by  mesquite  bushes  or  on  small 
dead  plants  above  the  water. 

June  15,  Beeville,  Tc\.,  at  night.  Acns  began  first,  soon  to  be  joined  by 
R.  pipiens,  then  came  M,  cincrca,  and  finally  one  or  two  Microhyla  ohvacea. 
,  .  .  We  caught  several  //.  cinerea.  In  one  mesquite  or  papilionaceous  plant 
found  t\vo  males  facing  each  other,  and  caught  each  by  putting  the  light  be- 
tween my  legs,  and  grabbing  with  each  hand.  //.  cinerea  were  about  I  or  2 
feet  above  the  \\atcr.  A  beautiful  brown  garter  snake  (Thamnophis  eqttes) 
was  coursing  around  the  pond,  ostensibly  for  frogs. 

June  17-21,  Brownsville-San  Benito,  Tex.  In  various  rcsacas  were  several 
choruses  of  H.  ctnerea. 


TREE  FROGS:  HYLIDAE 


307 


Plate  LXIV.  Hylacinerea  cinerea  (X%). 
3.  Females.  2.  Male. 


Plate  LXV.  Hyla  cinerea  cvittata.  1,3. 
Males  (Xi%).  2,  Female  (XO- 


308  HANDBOOK  OF  FROGS  AND  TOADS 

Authorities9  corner: 

W.  Bartram,  Gen.,  1791,  p.  277.  E.  A.  Andrews,  N.C.,  1928,  pp.  269-270. 

P.  H.  Pope,  Tex.,  1919,  pp.  95-96.  J.  D.  Kilby,  Fla.,  1945,  pp.  81-82. 

V.  R.  Haber,  Gen.,  1924,  pp.  1-32.  G.  L.  Orton,  La.,  1947,  pp.  364-369. 

Miller's  Tree  Frog,  Green  Tree  Frog 
Hyla  cinerea  evittata  (Miller).  Plate  LXV;  Map  22. 

Range:  Virginia  to  Maryland. 

"The  series  cvittata  in  the  American  Museum  differ  from  these  cinerea  in 
having  a  more  vertical,  less  sloping  profile  to  the  snout.  The  former  race  is 
also  said  to  differ  from  cinerea  m  its  broader  head  but  our  series  of  ctnerea  ex- 
hibits a  great  variation  in  width  and  no  constant  difference  can  be  found  here. 
It  thus  appears  that  typical  cinerea  has  a  more  northern  range  than  Wright 
and  Wright  (1933)  have  assumed.  It  follows  that  cvittata  has  a  more  restricted 
range  for  we  found  only  cinerea  m  the  Cove  Point-Solomon's  Island  area.  It 
would  be  interesting  to  know  if  all  green  frogs  north  of  the  Washington  area 
are  not  referable  to  cinerea"  (G.  K.  Noble  and  W.  G.  Hassler,  Md.,  1936, 

P-fy)« 
Since  1912  we  have  been  seeing  evittatc,  semifasciate,  and  striped  cinerea 

from  Virginia  to  Bay  St.  Louis,  Miss.,  and  did  not  assume  as  intimated  above. 
We  merely  by  our  action  provisionally  called  the  northern  population  (varia- 
ble though  it  is  even  at  Mt.  Vcrnon  where  we  have  collected  it)  evittata.  Col- 
lectors fiom  West  Virginia  to  Florida  to  Louisiana  have  recorded  or  collected 
evittatc  cinerea.  It  is  interesting  to  see  that  E.  R.  Dunn  (Va.,  1937,  p.  10)  so 
views  the  northern  collection  of  material: 

"To  sum  up:  ST  per  cent  in  the  upper  tidewater  Potomac  area  have  no  stripe 
or  a  short  stripe;  41  per  cent  in  other  parts  of  Maryland  and  Virginia  have 
no  stripe  or  a  short  one.  Carolina  material  available  to  me  is  not  very  extensive, 
but  it  would  seem  that  there  only  25  per  cent  have  no  stripe  or  a  short  stripe, 
whereas  7*5  per  cent  have  a  long  stripe.  Reports  from  further  south  indicate 
that  TOO  per  cent  long  stripe  occurs  in  the  far  south,  especially  on  the  Gulf 
("oast. 

"We  arc,  therefore,  faced  with  two  opposed  populations,  obviously  differ- 
ent. One  occurs  in  the  upper  tidewater  Potomac;  the  other  occurs  in  the  far 
south.  An  intermediate  population  occurs  over  a  wide  area.  Unfortunately  a 
somewhat  intermediate  population,  that  of  the  Carolinas,  was  named  first. 
This  seems  to  be  nearer  that  of  the  far  south,  so  that  Hyla  cinerea  cinerea  may 
be  properly  applied  to  specimens  of  Hyla  cinerea  from  the  Carolmas  south. 
The  name  Hyla  cinerea  evittata  may  be  properly  applied  to  the  upper  tide- 
water Potomac  population.  The  rest  of  the  Maryland  and  Virginia  popu- 
lations are,  and  should  be  considered,  intermediate  between  cinerea  and 
evittata. 


TREE  FROGS:  HYLIDAE  309 

"The  most  northern  locality  yet  known  is  the  western  end  of  the  Chesa- 
peake-Delaware canal  in  Cecil  Co.,  Md.,  reported  to  me  by  Mr.  Joseph 
Cadbury.  It  is  unknown  from  Delaware  or  from  the  eastern  side  of  the  Del- 
Mar-Va  peninsula.'* 

Habitat:  These  frogs  live  in  lily  pools  and  reed  beds  of  tid^l  marshes.  At 
other  times,  they  are  found  on  bushes  and  small  trees  near  the  water. 

"Very  little  is  known  about  the  habits  of  Hyla  evittata.  In  June  and  July  the 
animals  are  to  be  found  in  the  rank  vegetation  of  the  tide  marshes.  Here  they 
remain  quiet  during  the  day,  but  as  evening  approaches  they  become  active 
and  noisy.  Their  food  at  this  time  consists  chiefly  of  a  small  beetle  that  is  found 
on  the  leaves  of  the  pond-lilies.  ,  .  .  Later  in  the  season  the  frogs  leave  the 
low  marsh  vegetation.  As  they  are  then  perfectly  silent  they  are  difficult  to 
find,  though  occasionally  one  may  be  seen  in  a  bush  or  small  tree,  but  never 
far  from  water"  (G.  S.  Miller,  Jr.,  D.C.,  iHw,  p.  78). 

Recently  (Va.,  1944,  p.  1187)  Dr.  P.  Bartsch  wrote:  "I  have  been  rather  inter- 
ested in  the  habits  of  Hyla  evittata  Miller,  as  observed  on  our  farm,  which 
joins  Pohick  Bay,  a  tributary  of  the  Potomac  River,  some  21  miles  south  of 
Washington.  Our  house  is  approximately  two-fifths  of  a  mile  from  the  water 
at  an  elevation  of  155  feet. 

"In  the  late  summer  when  the  young  hylas  have  reached  a  little  more  than 
half  their  adult  size  they  leave  the  lily  pads  and  bonnets  of  the  bay  and  move 
inland.  At  this  time  the  frogs  come  to  our  upland  level,  where  we  find  them 
particularly  partial  to  okni  plants.  I  have  found  as  many  as  six  attached  to  the 
stems  of  a  short  row  of  this  vegetable;  they  blend  beautifully  into  the  okra 
color  scheme. 

"All  the  specimens  caught  on  our  place  arc  evittata;  that  is,  they  lack  the 
band  characteristic  of  H.  ctnerea.  On  the  north  side  of  the  Potomac  River  at 
Foxes  Ferry  I  have  found  the  striped  form  predominant  and  the  evittata  color 
scheme  only  slightly  represented." 

Size:  Adults,  I'/j-i^s  inches.  Males,  36-47  mm.  Females,  32-47  mm. 

General  appearance:  This  slim,  smooth,  green  irog  is  like  Hyla  cincrea,  but 
without  the  light  stripe  on  the  sides  and  legs.  The  under  parts  are  white, 
ivory  yellow,  or  maigucritc  yellow,  with  purple-vmaccous  to  brownish  vina- 
ccous  on  the  front  of  the  foicarms,  femur,  the  rear  of  the  tibia,  and  more  or 
less  on  the  underside  of  the  legs.  The  rear  of  arms,  legs,  and  hind  feet  arc  mar- 
gined with  white  to  marguerite  yellow.  Sometimes  there  arc  fine  yellow  spots 
on  the  back. 

Color:  Male.  Upper  parts  cosse  green  or  lettuce  green;  sides  and  legs  javel 
green  or  dull  green-yellow;  sides  of  under  jaw  from  angle  almost  to  tip  nar- 
rowly bordered  with  apple  green  or  dull  green-yellow;  rear  of  arms,  legs,  and 
hind  feet  margined  with  white  to  marguerite  yellow;  front  of  legs,  feet,  axilla, 
and  somewhat  on  underside  of  forearm  purple-vinaceous,  livid  brown,  or 
deep  brownish  vinaceous;  under  parts  including  throat  white,  ivory  yellow, 


310  HANDBOOK  OF  FROGS  AND  TOADS 

or  marguerite  yellow.  This  individual  had  faint  line  on  upper  jaw  tip  to  angle 
of  mouth  primrose  yellow. 

Female.  This  one  has  the  faintest  line  on  upper  jaw.  Upper  parts  oil  green; 
oil  green  on  sides  of  under  jaw  as  prominent  as  in  male;  under  parts  white  or 
cartridge  buff;  front  of  forearms,  femur,  rear  of  tibia,  underside  of  tarsus  and 
foot  deep  brownish  vinaceous.  Another  female  has  entire  under  parts  of  legs 
and  most  of  pectoral  region  purplish  vinaceous;  line  on  upper  jaw  white;  fine 
lemon  yellow  or  light  cadmium  spots  on  the  back. 

Structure:  Broader  head  and  deeper  face  than  Hyla  anerea. 

"Hyla  cvittata  sp.  nov.  Type  adult  &  (in  alcohol)  No.  26,291  U.S.N.M., 
collected  at  Four  Mile  Run,  Alexandria  County,  Virginia,  July  15,  1898,  by 
Gcrrit  S.  Miller,  Jr.,  and  Edward  A.  Prcble.  Zonal  position. — This  frog  is 
probably  confined  to  the  Upper  Austral  zone.  Geographic  distribution. — 
While  the  animal  is  at  present  known  from  the  marshes  of  the  Potomac  River 
near  Washington  only,  it  is  to  be  looked  for  near  the  coast  from  Chesapeake 
Bay  to  Long  Island  Sound.  General  characters. — Like  Hyla  cinerea  (Daudin) 
but  with  broader,  deeper  muzzle  and  normally  unstriped  body  and  legs. 
Color. — Entire  dorsal  surface  varying  from  olivaceous  brown  through  deep 
myrtle-green  to  pale  yellowish  grass-green;  ventral  surface  white,  irregularly 
tinged  with  yellow,  especially  on  chin  and  throat;  colors  of  back  and  belly 
fading  rather  abruptly  into  each  other  on  lower  part  of  sides;  skin  of  under 
surface  of  limbs  unpigmcntcd,  transparent;  legs  and  jaws  slightly  paler  on 
sides  than  above;  eye  very  bright  and  iridescent,  the  pupil  black,  the  iris 
golden  greenish  yellow,  thickly  dotted  with  black,  back  with  a  few — usually 
less  than  half  a  do/en — inconspicuous  minute,  yellowish  dots.  Measurements. 
— Type:  head  and  body,  48;  hind  leg,  6y;  femur,  20;  tibia,  21;  tarsus,  it;  hind 
foot,  17;  humcrus,  8;  forearm,  8;  front  foot,  10;  greatest  width  of  head,  14; 
eye  to  nostril,  $.5;  distance  between  nostrils,  3.5.  An  adult  cf  from  the  type 
locality:  head  and  body,  s<>;  hind  leg,  70;  femur,  21;  tibia,  21;  tarsus,  ri;  hind 
foot,  17;  humeius,  8;  forearm,  8;  front  foot,  10;  greatest  width  of  head,  14;  eye 
to  nostril,  4;  distance  between  nostrils,  3"  (G.  S.  Miller,  Jr.,  D.C.  and  Va.,  1899, 
p.  76) i . 

Voice:  The  call  is  very  like  that  of  Hyla  cinerea. 

"The  note  is  like  that  of  Hyla  ptcl(eringn  in  form,  but  in  quality  it  is  com- 
paratively harsh  and  reedy,  with  a  suggestion  of  distant  guinea-fowl  chatter, 
and  scarcely  a  trace  of  the  peculiar  freshness  so  characteristic  of  the  song  of 
the  smaller  species.  The  song  period  continues  through  June  and  July"  (G.  S. 
Miller,  Jr.,  Va.,  181)9,  p.  78). 

Breeding:  Similar  to  H.  cinerea.  Smallest  specimens  in  National  Museum 
February,  1928,  were  two  young  taken  by  A.  S.  Miller  at  Quantico,  Va.,  Oct. 
1 5,  igoi.  They  were  22-24  mm-  Another  of  26  mm.  was  taken  September,  1923, 
at  New  Alexandria,  Va.,  by  Dr,  E.  T.  Wherry.  These  are  5-15  mm.  larger 
than  the  11-17  mm.  transformces  of  H.  cinerea  cinerea. 


TREE  FROGS:  HYLIDAE  3n 

Journal  notes:  In  1917,  on  May  29,  while  encamped  on  the  Alexandria 
(Henshaw  Sparrow)  Heights,  we  heard  Hyla  evittata  toward  the  river.  In 
1922,  in  June,  we  heard  them  again  from  Alexandria.  One  member  of  the 
party  who  knew  and  had  collected  H.  c.  evittata  thus  identified  them.  The 
calls  impressed  the  authors  as  Hyla  cinerea.  On  June  3,  1928,  Mr.  and  Mrs. 
Morris  Brady  took  us  to  one  of  the  best  collecting  grounds  of  Hyla  c.  evittata. 
We  walked  along  the  railroad  track  and  picked  up  one  or  two  specimens  of 
Hyla  evittata  on  the  ties,  as  well  as  a  few  toads.  On  the  right-hand  side  of  the 
track,  Brady  took  me  to  the  water's  edge  where  we  heard  a  few  calling.  It  cer- 
tainly sounded  like  Hyla  c.  cinerea.  Captured  one  more.  Luckily  for  me,  both 
sexes  are  represented.  These  specimens  are  evittate,  but  it  would  not  be  hard 
to  conceive  of  some  of  them  developing  a  complete  stripe  or  a  semifasciate  con- 
dition. In  fact,  Mr.  Brady  informs  me  that  individual  frogs  may  take  on  these 
different  liveries.  The  life  history  of  this  form,  according  to  what  he  told  me, 
is  very  similar  to  that  of  Hyla  c.  cinerea.  We  await  with  expectancy  his  account 
of  this  species. 

Authorities9  corner: 
G.  S.  Miller,  D.C.  and  Va.,  1899,  pp.  75-     G.  S.  Myers,  N.C.,  1924,  p.  60. 

78.  O.  C.  Van  Hyning,  Fla.,  1933,  p.  4. 

W.  P.  Hay,  D.C.,  1902,  p.  199.  N.  D.  Richmond  and  C.  J.  Coin,  Va., 

H.  W.  Fowler,  Md.,  1915,  pp.  38-39.  J938,  p.  303. 

"The  material  at  hand  indicates  that  the  population  of  this  frog  inhabiting 
the  Del-Mar-Va  Peninsula  is  intermediate  between  the  two  subspecies,  Hyla 
cinerea  cinerea  (Schneider)  and  Hyla  anerea  evittata  Miller"  (R.  Conant, 
Del.,  1945,  p.  4). 

Peeper,  Spring  Peeper,  Pickering's  Tree  Frog,  Pickering's  Tree  Toad,  Pick- 
ering's Hylodes,  Pickering's  Hyla,  Peeping  Frog,  Castanet  Tree  Frog 

Hyla  cmcifer  crucifer  Wied.  Plate  LXVI;  Map  23. 

Range:  Gaspe  Peninsula  to  Manitoba  and  Minnesota  south  to  northeast- 
ern Kansas,  Arkansas,  Louisiana,  Texas,  Mississippi,  Alabama,  Piedmont 
Georgia,  South  Carolina  to  New  Brunswick. 

Habitat:  These  frogs  live  in  open  lowland  marshes,  swamps  at  sources  of 
streams  whether  wooded  or  open,  sphagnous  or  cattail,  in  fact  any  pool,  ditch, 
or  shallow  pond  transient  or  permanent,  grassy  or  muddy. 

Size:  Adults,  %-iM  inches.  Males,  18-29  mm.  Females,  20-^3  mm. 

General  appearance:  This  is  a  small  frog  with  an  oblique  cross  on  the  back. 
A  male  when  first  captured  in  early  spring  may  be  liver  brown  or  bay  to  claret 
brown.  The  females  are  usually  lighter  in  color.  Both  have  obscure  bands 
across  the  fore  and  hind  limbs.  The  male  has  yellow  on  the  groin;  its  throat 
is  olive.  The  under  parts  are  light  vinaceous-cinnamon  to  pale  yellow. 

Color:  M ale.  Upper  parts  deep  olive  buflf  or  Isabella  color  with  light  brown- 


3I2 


HANDBOOK  OF  FROGS  AND  TOADS 


ish  olive  on  back  and  obscure  bands  across  fore  and  hind  limbs;  primrose  yel- 
low on  groin;  seafoam  green  or  glass  green  later  ad  of  vent;  below  vent,  natal 
brown  or  clove  brown  spot;  throat  olive-ocher,  or  aniline  yellow  in  rear  and 
citrine  near  lower  jaw  rim;  pectoral  region  sulphur  yellow  or  pale  chalcedony 


Map  23 

yellow;  rest  of  under  parts  light  vinaceous-cinnamon;  upper  jaw  cinnamon- 
buff;  bister  stripe  in  front  of  eye.  A  male  when  first  captured  out  in  the  sun  in 
early  spring  may  be  liver  brown,  chestnut  brown,  bay,  claret  brown,  or  mars 
brown. 


TREE  FROGS:  HYLIDAE  313 

Female.  Upper  parts  warm  buff,  cinnamon-buff,  cream-buff,  chamois.  Bar 
between  the  eyes,  cross  on  fore  part  of  back,  oblique  bar  back  of  it  on  left  and 
also  right,  bar  on  back  near  vent,  bars  on  forearms  and  also  on  tibia  buffy 
brown  or  drab.  Stripe  in  front  of  eye  and  across  tympanum  and  spot  on  vent 
olive  brown  or  hair  brown.  Venter  pale  olive-buff,  cartridge  buff,  or  cream 
color.  Rear  of  hind  legs  and  front  of  thigh  aniline  yellow,  olive-ocher.  Under- 
side of  femur  vinaceous-fawn  to  light  grayish  vinaceous.  Iris  vinaceous-cinna- 
mon  or  onion-skin  pink  with  some  dark  spots. 

Structure:  Muzzle  pointed,  projecting  considerably  beyond  the  lower  jaw; 
skin  smooth  or  nearly  so;  fingers  not  webbed. 

Voice:  Notes  are  variable,  but  shrill,  clear,  high  pitched,  and  can  be  heard 
at  a  distance  of  half  a  mile.  When  at  the  crest  of  the  chorus  stage,  the  din  may 
be  heard  incessantly  night  and  day,  though  it  is  most  vociferous  in  the  after- 
noon and  evening.  The  male  approaching  the  pond  gives  a  weaker  note  with 
a  more  querulous  tremor  in  it. 

"The  vocal  sac  of  the  male  is  largely  developed  in  spring;  it  is  of  a  greenish 
grey  color  and  lies  in  loose  folds  outside  the  muscles  of  the  throat.  Inside  the 
mouth  are  two  slits  or  orifices,  opening  into  the  sac,  one  on  each  side  near 
the  angle  of  the  jaws.  When  the  frog  is  about  to  give  voice  the  whole  body  is 
inflated,  followed  by  that  of  the  vocal  sac  which  rounds  out  into  a  bubble  and 
does  not  collapse  with  each  'peep';  the  degree  of  inflation  evidently  governs 
the  volume  of  sound.  About  sunset  I  have  frequently  been  aided  in  discover- 
ing the  frogs  by  the  level  rays  of  the  sun  striking  along  the  edge  of  pond  or 
meadow  and  reflecting  from  this  moist,  inflated  vocal  sac  with  a  glittering 
light. 

"Unless  the  day  is  overcast,  or  a  warm  rain  is  falling,  little  is  to  be  heard 
from  the  frogs  till  about  four  o'clock  in  the  afternoon  when  their  concerts 
begin,  to  be  continued  in  mild  nights  till  morning.  Considering  the  size,  the 
volume  of  sound  possible  from  one  frog  is  surprising.  As  you  approach  a 
locality  where  they  are  in  full  voice  the  air  seems  to  grow  gradually  dense 
with  this  ear-deafening,  all-pervading  sound;  occasionally  the  voices  fall  into 
a  regular  measure  of  time,  but  the  effect  is  usually  a  medley  of  shrill  sounds, 
a  few  voices  audible  above  the  others  by  reason  of  some  peculiarity  in  key, 
or  lack  of  smoothness  in  utterance.  The  piping  of  each  individual  is  long  con- 
tinued; the  interval  between  these  musical  efforts  appears  to  depend  on  the 
mood  of  the  musician.  One  does  not  note  the  pause  of  individual  voices  in 
the  general  effect,  but,  however  loud  and  earnest  the  piping  may  be,  the  in- 
troduction of  any  unusual  sound  or  appearance,  even  the  low  quick  flight  of 
a  bird  over  the  water,  is  almost  sure  to  give  alarm  and  still  them  for  a  while; 
the  frogs  along  the  edge  of  the  shore  commonly  settling  away  out  of  sight  for 
safety  among  the  dead  leaves  under  water,  while  those  having  a  position  on 
the  low  bushes  or  reeds  merely  cling  more  closely,  flattening  the  body  against 
the  object  on  which  they  are  resting.  The  interval  of  silence  is  brief;  soon  a 


314  HANDBOOK  OF  FROGS  AND  TOADS 

frog  rises  and  gives  a  shrill  'peeP>*  which  is  immediately  answered  by  dozens 
of  voices.  The  sounds  may  appear  to  come  from  about  your  feet,  but  for  the 
reasons  given,  the  chances  are  against  seeing  the  frogs  till  some  movement  in 
the  water  as  they  rise  from  their  hiding  places,  arrests  the  eye,  which  on  per- 
ceiving one,  usually  discovers  more"  (M.  H.  Hinckley,  Mass.,  1883,  pp.  314- 


Breeding:  They  breed  from  April  i  to  June  15.  The  eggs,  singly  laid,  arc 
submerged  among  fine  grass  or  other  plants  in  matted  vegetation,  usually  near 
the  bottom  of  the  pond,  and  are  800-1000  in  number.  They  are  white  or 
creamy  and  black  or  brownish  in  color,  the  jelly  firm  with  well-defined  out- 
line; the  egg  K'5  inch  (0.9-1.1  mm.),  the  envelope  %o-/42  inch  (1.2-2.0 
mm.).  The  small  tadpole,  i%  inches  (33  mm.),  has  tooth  ridges  %  or  %. 
After  90  to  100  days  the  tadpoles  transform  from  July  i  to  Aug.  i,  at  %-%6 
inch  (9-14  mm.). 

Journal  notes:  It  is  interesting  to  observe  how  suddenly  a  chorus  will  end 
at  one's  approach,  only  to  be  resumed  if  the  intruder  remains  quiet.  At  this 
time,  the  frogs  may  be  among  the  grassy  hummocks  along  the  sides  of  the 
ponds,  or  in  the  shallow  pools  within  the  surface  film  of  dead  leaves  and 
algae.  Not  infrequently  when  disturbed,  they  may  be  seen  leaping  on  this 
matted  carpet  before  disappearance.  In  one  instance  we  discovered  one  in  the 
spathe  of  the  skunk  cabbage. 

March  29,  1910.  On  the  10  A.M.  trip,  noticed  a  pair  of  peepers  come  to  the 
surface.  Watched  them  return  to  the  bottom.  The  male,  as  always  before,  was 
almost  as  dark  as  the  darkest  dead  leaf  of  spring  (in  the  pond)  ;  the  female 
was  much  lighter,  about  as  light  as  both  males  and  females  are  at  night.  The 
female  would  lay  an  egg,  then  walk  or  drag  along  3  or  4  steps,  then  stop  for 
another  fertilization  and  emission;  and  so  they  went  on  some  time  through 
weeds  and  matted  vegetation  without  rising  to  the  surface  during  my  observa- 
tions. Probably  they  came  up  occasionally,  as  when  I  first  discovered  them. 
Put  them  in  a  wet  handkerchief  and  then  in  a  glass  jar  where  the  sun's  rays 
could  not  affect  them;  took  them  to  a  crossroads  pond.  When  I  planted  the 
jar,  they  were  both  lighter  in  color  than  when  captured,  the  male  considerably 
so.  As  I  put  the  jar  in  the  pond,  they  broke  embrace  but  resumed  later  and 
laid  eggs  later  in  the  day. 

April  7,  1929.  Went  to  Ringwood  (Ithaca,  N.Y.)  for  peepers  and  wood 
frogs.  Went  about  6  o'clock.  As  it  grew  dusky,  the  dm  grew  louder.  Peepers 
seemed  all  around  us,  in  ponds  and  on  land,  on  the  ground  and  on  bushes. 
The  calls  were  everywhere,  but  to  find  one  and  see  him  call  was  a  real  job. 
We  both  looked  for  some  time  without  success.  Then  we  both  concentrated 
on  one  small  pond  near  the  road  that  had  cattails  in  it,  also  a  fence  post,  a 
fallen  log  or  rail,  and  a  few  bushes  around  the  edge.  It  was  largely  made  up 
of  tussocks  of  grass  or  sedge  standing  in  water.  The  field  grass  at  one  edge 
was  wet  and  "oozy."  We  knew  we  must  be  looking  directly  at  several,  but 


TREE  FROGS:  HYLIDAE 


315 


there  was  so  much  noise  it  was  hard  to  single  out  one  note  and  locate  it. 
Finally  I  saw  one.  It  was  down  in  the  water,  just  at  the  surface,  standing  al- 
most erect,  sunken  into  the  edge  of  a  tussock,  his  back  toward  the  water,  the 
throat  bubble  toward  the  grass  stems.  He  looked  dark  greenish  brown  or 
almost  black,  like  the  dirty  bases  of  the  grass  stems.  If  his  bubble  hadn't  been 
vibrating,  I  never  would  have  seen  him.  At  the  same  instant,  Bert  caught  one. 
Then  we  found  a  pair,  the  female  looking  much  lighter.  These  were  at  a 
central  tussock.  We  found  another  pair  at  the  edge  in  the  "sopping"  field 
grass.  Then  several  feet  away  from  the  pond,  three  feet  up  on  a  weed  stem, 
was  a  male  calling  lustily.  Then  we  went  over  to  the  ponds  in  the  woods.  Here 
were  peepers  everywhere  m  ponds  and  on  land.  One  little  fellow  was  cling- 
ing to  a  stem  a  foot  or  so  from  the  ground,  clinging  rather  diagonally  so  that 
his  bubble  went  to  one  side  of  the  stem.  We  counted  his  call.  It  seemed  to  be 
about  one  a  second,  fifteen  calls  or  so,  and  then  a  pause.  He  sat  for  periods  with 
his  throat  extended.  On  a  log  extending  into  the  water,  Bert  saw  six  peepers 
calling,  but  he  slipped  from  the  log  and  they  stopped. 

July  14,  1929.  Went  to  Ringwood  for  adult  wood  frogs.  Along  the  wood 
road  in  a  fairly  open  spot  where  berry  bushes  and  grass  are  creeping  in  found 
many  transformed  peepers.  Some  were  on  the  ground  in  an  area  covered 
sparsely  with  a  fine,  soft  grass.  Many  were  sitting  on  the  leaves  of  the  berry 
bushes  2  or  3  feet  above  the  ground,  others  were  hopping  over  the  dead  leaves 
in  the  woods.  In  the  pond,  along  with  the  numerous  newt  adults  and  larvae, 
countless  larvae  of  the  spotted  salamander,  a  few  transforming  pickerel  frogs, 
and  a  very  few  transforming  wood  frogs,  were  many  peepers  in  various  stages 
of  transformation. 

August  2,  1929.  In  the  same  woods,  many  transformed  and  a  few  adult 
peepers  were  on  the  forest  floor.  In  the  pond,  we  found  no  mature  peeper  tad- 
poles, and  only  three  or  four  transforming  ones. 

Authorities'  corner: 
J.  A.  Allen,  Mass.,  1868,  p.  190. 
M.  H.  Hinckley,  Mass.,  1883,  pp.  313-317. 
W.  I.  Sherwood,  Conn.,  1898,  p.  19. 

Florida  Peeper,  Southern  Peeper,  Bartramian  Peeper,  Southern  Spring 
Peeper,  Sabalian  Peeper,  Coastal  Peeper 

Hyla  cruet jer  bartramtana  Harper.  Plate  LXVII;  Map  23. 

Range:  Coastal  Plain  of  southern  Georgia  and  northern  Florida.  Doubtless 
some  of  the  coastal  records  of  Alabama  and  Mississippi  are  of  this  form. 

Habitat:  Similar  to  that  of  H.  c.  cructfer.  "Ponds  and  ditches  in  and  near 
woods;  moderately  common.  Begins  to  call  in  December  and  January,  on 
warm  nights"  (O.  C.  Van  Hymng,  Fla.,  1933,  p.  4). 

"The  six  specimens  from  Alachua  County,  Florida,  listed  by  Harper  .  ,  , 


3i6 


HANDBOOK  OF  FROGS  AND  TOADS 


Plate  LXVI.  Hyla  crucijcr  crttcifer.  1,2,3.  Plate  LXV1I.  Hyla  crucifer  bartramiana 

Males  (XO-  4-  E8gs  (X1^)-  5-  Female          (XO- 


TREE  FROGS:  HYLIDAE 

in  the  original  description  of  this  form,  were  taken  in  a  small  woods  pond 
which  is  nearly  completely  covered  with  hyacinths.  The  greatest  number  of 
individuals  I  have  seen  at  any  one  time  in  northern  Florida  was  in  the  above 
mentioned  pond,  where  I  collected  eighteen  on  the  evening  of  December  31, 
1940"  (C.  J.  Coin,  Fla.,  1943,  p.  149). 

"Northern  Florida  except  in  the  western  portion  of  the  Panhandle.  Known 
from  the  following  counties:  Jackson,  Liberty,  Baker,  Alachua,  Citrus  and 
Lake.  Habitat.— Mesophytic  and  low  hammock,  swamp  borders,  the  more 
open  bay-heads,  and  tangles  along  the  smaller  streams.  Abundance.— Moder- 
ately common  during  breeding  season;  difficult  to  collect  in  numbers  during 
summer  and  fall.  Habits.— They  hibernate  and  aestivate  under  logs  and  bark 
and  in  knot-holes.  In  Alachua  County  they  occupy  essentially  the  same  habi- 
tats as  the  much  rarer  H.  v.  versicolor"  (A.  F.  Carr,  Jr.,  Fla.,  1940,  p.  59). 

Size:  Harper's  type,  a  male,  is  28  mm.;  the  female  topotypes  are  31.0  and 
33.5  mm. 

General  appearance:  "Six  specimens  of  bartramiana  from  Alachua  County, 
Florida,  generously  presented  by  Coleman  J.  Coin,  exhibit  a  still  more  distinct 
spotting  of  the  under  parts  than  specimens  from  southern  Georgia  and  proba- 
bly represent  the  extreme  in  subspecific  characters"  (F.  Harper,  Ga.,  i93Qb, 

P.  2). 

"Aside  from  the  average  difference  in  spotting,  the  Coastal  Plain  specimens 
were  noticeably  larger  than  those  of  the  Piedmont"  (F.  Harper,  Ga.,  1939!), 

P- 3)- 

Color:  Gainesville,  Fla.,  from  Coleman  J.  Coin,  Feb.  16,  1946.  Descriptions 
of  several  widely  varying  patterns  follow : 

1.  Back  orange-cinnamon  with  bars  on  the  back  of  snuff  brown.  Of  the 
bars,  there  are  2  prongs  of  the  cross  toward  eyes  and  2  down  sides  back  of  mid- 
body;  between  lateral  bars  of  cross  there  may  be  intermediate  series  connected 
or  not;  in  front  of  insertion  of  hind  legs  is  an  oblique  bar  extending  up  onto 
the  back;  also  a  bar  between  eyes  going  onto  eyelid;  the  same  color  goes  from 
nostril  through  eye  and  halfway  along  side.  Color  of  groin,  front  and  rear  of 
femur,  and  lower  side  of  tibia  is  yellow  ocher  to  buckthorn  brown;  rear  of 
femur  heavily  spotted  amber  brown.  Upper  side  of  fore  and  hind  legs  orange- 
cinnamon  with  prominent  transverse  bars  on  femur  and  tibia,  line  on  tarsus 
and  foot  of  claret  brown  bordered  above  by  thread  of  capucine  buff.  Venter, 
capucine  orange  to  light  ochraceous  salmon  heavily  spotted  on  breast  with 
amber  brown;  throat  light  orange-yellow  in  mid-portion  with  rim  of  lower 
jaw  black.  Iris,  lower  part  snuff  brown  like  vitta,  upper  part  mikado  orange. 

2.  Back  xanthine  orange,  groin  cadmium  yellow  or  raw  sienna.  No  black 
on  throat. 

3.  Back  cinnamon. 

4.  Back  buffy  brown,  groin  bar  goes  completely  across  the  back.  Rear  of 


3i8  HANDBOOK  OF  FROGS  AND  TOADS 

femur  extensively  spotted  and  bars  of  tibia  extend  onto  orange  area.  Center 
of  throat  olive  lake,  shading  to  black  on  margin  of  lower  jaw. 

5.  Back  pinkish  cinnamon,  bars  buffy  olive.  Venter  amber  yellow,  groin 
and  rear  of  femur  capucme  yellow. 

6.  Back  Isabella  color  with  face  mask  and  bars  of  buffy  olive;  line  along 
tarsus  madder  brown;  throat  Saccardo's  olive  with  madder  brown  flecks. 

7.  Back  mummy  brown,  no  bars  showing;  bars  on  legs  black;  dorsum  of 
legs  buckthorn  brown;  orange  groin  and  both  sides  of  femur;  line  along  tar- 
sus claret  brown;  throat  Saccardo's  olive  to  black  on  margin  of  jaw;  belly 
cinnamon-buff. 

8.  Back  ochraceous-tawny  with  Saccardo's  olive  bars. 

9.  Back  kaiser  brown  with  markings  of  carob  brown. 

Remarks:  March  29,  1946.  Live  material  from  Gainesville,  Fla.,  and  Ithaca, 
N.Y.,  compared.  Differences  most  apparent  are:  the  Florida  frogs  are  deeper 
orange  on  the  belly,  with  more  dark  specks  on  under  parts  and  with  the  back 
of  the  pecan  brown  order,  whereas  those  of  Ithaca  are  tawny-olive.  When  we 
first  glanced  at  the  Florida  forms  we  concluded  that  there  were  more  acces- 
sory arms  to  the  dorsal  cross  than  in  northern  forms,  i.e.,  6  or  7  arms  instead 
of  4,  but  some  northern  ones  have  the  accessory  branches  also.  In  general,  the 
cross  and  bars  of  the  Florida  form  are  somewhat  broader  and  more  conspicu- 
ous, the  bars  on  legs  of  some  northern  forms  being  very  narrow.  The  belly  of 
northern  ones  is  vinaceous,  whereas  Florida  frogs  are  nearer  honey  yellow, 
a  color  strikingly  apparent  on  concealed  surfaces  of  hind  legs  and  underside 
of  femur.  Furthermore  the  rear  of  femur  in  most  of  the  Florida  frogs  has 
many  fine  black  specks  that  are  lacking  in  our  Ithaca  ones. 

Structure:  "Length  (snout  to  vent),  28  mm.;  tibia,  14;  whole  hind  foot, 
19.5;  elbow  to  tip  of  third  finger,  13;  interolecranal  extent  (distance  between 
elbows  when  humeri  are  extended  in  the  same  line  at  right  angles  to  longi- 
tudinal axis  of  body),  17.5;  intergenual  extent  (distance  between  knees  when 
femora  are  extended  in  the  &ame  line  at  right  angles  to  longitudinal  axis  of 
body),  26"  (F.  Harper,  Ga.,  1939^  pp.  1-2). 

Voice:  "The  males  sing  in  buttonbush,  briars,  willows,  Decodon,  etc.,  at  the 
water's  edge  in  small  ponds,  ditches  and  flooded  meadows"  (A.  F.  Carr,  Jr., 
Fla.,  1940,  pp.  59-60). 

(See  Psetidacris  ornata  for  Carr's  1933  comparisons  of  calls  of  P.  ornata  and 
H.  c.  bartranriana.} 

Breeding:  "In  Charlton  County,  Georgia,  for  example,  the  breeding  season, 
as  far  as  may  be  judged  by  vigorous  choruses,  extends  from  late  November 
more  or  less  continuously  to  early  March.  Eggs  were  collected  December  9. 
...  All  (tadpoles)  lack  the  purplish  black  blotches  along  the  outer  edge  of 
the  tail  crests  which  Wright  records  .  .  ."  (F.  Harper,  Ga.,  1939,  p.  3). 

"January  i  to  March  24.  Apparently  the  season  is  much  earlier  in  Florida 


TREE  FROGS:  HYLIDAE  319 

than  farther  north,  where  it  is  April  i  to  June  15  according  to  Wright.  .  .  . 
I  have  found  them  breeding  when  the  temperature  was  35°  F.  P.  ornata  is  the 
only  other  Florida  frog  which  breeds  during  such  cold  weather"  (A.  F.  Carr, 
Fla.,  1940,  pp.  59-60). 

"In  the  Gainesville  region  the  peepers  usually  enter  the  ponds  and  marshes 
in  late  December,  and  continue  to  call  spasmodically  until  early  spring.  My 
only  collection  records  are  for  February  and  December"  (C.  J.  Coin,  Fla., 
1943,  p.  149). 

Journal  notes:  We  have  had  no  experience  with  this  form  in  the  field. 

Authorities'  corner: 
A.  F.  Carr,  Jr.,  Fla.,  1940,  p.  59. 

Pine  Woods  Tree  Frog,  Pine  Woods  Tree  Toad,  Pine  Tree  Toad,  Pine  Tree 
Frog,  Scraper  Frog,  Femoral  Hyla 

Hyla  femoraKs  Latrcillc.  Plate  LXVIII;  Map  21. 

Range:  Coastal  region  o£  Maryland  to  Florida  to  Louisiana.  We  rather 
question  trans-Mississippian  records  to  Dallas,  Tex.;  they  are  possible  but 
doubtful.  Like  Hyla  gratiosa,  at  times  H.  femoralis  betakes  itself  into  the  high 
pines  and  is  hard  to  get,  and,  also  like  H.  gratiosa,  it  is  a  decidedly  Sabalian  or 
Gulf  Strip  species. 

Habitat:  Trees  and  shrubs  of  pine  barrens.  It  breeds  in  grassy  transient  pools 
at  the  roadside  or  in  the  woods,  in  cypress  ponds  or  bays  or  in  lily-covered 
swamp  prairies. 

"High  pine,  high  hammock,  and  all  types  of  flatwoods,  occasionally  (at 
least)  climbing  to  the  tops  of  the  tallest  long-leaf  pines.  ...  I  have  found 
them  hibernating  in  rotten  pine  logs;  one  was  disinterred  at  a  depth  of  two 
feet  in  nearly  dry  sand,  Charlotte  County,  Dec.  16, 1934"  (A.  F.  Carr,  Jr.,  Fla., 
iQ4ob,  p.  60). 

"This  species  is  said  to  stay  high  up  in  pine  trees  but  the  only  two  I  have 
personally  taken  were  respectively  seven  feet  up  in  a  small  deciduous  tree, 
quite  likely  a  sweet  gum,  near  a  stream  in  the  first  case,  and  among  rank  vege- 
tation in  the  pine  barrens  in  the  second  case"  (C.  S.  Brimley,  N.C.,  1940,  p.  22). 

R.  F.  Deckert  in  1915  (Fla.,  p.  3)  said:  "Hyla  femoralis  Latreille  is  called 
the  Pine  Tree  toad,  from  its  habit  of  frequenting  the  tops  of  pine  trees  almost 
exclusively  during  the  summer  months." 

"In  1925  Viosca  in  discussing  the  Florida  Parishes  or  uplands  of  Louisiana 
comments  that  To  the  east  of  the  Shortleaf  Pine  area  are  the  Longleaf  Pine 
Hills  with  gentle  slopes,  and  intertwined  by  winding  creeks.  Here  we  have  a 
limited  representation  of  practically  all  species  found  in  the  uplands  generally 
and  in  addition,  some  species  which  may  be  said  to  be  characteristic.  Hyla 
femoralis  is  the  typical  tree  frog  of  this  section.' 


320  HANDBOOK  OF  FROGS  AND  TOADS 

"At  Leroy,  Ala.,  June  12,  1917,  Dr.  H.  H.  Knight  while  sweeping  bushes 
and  lower  branches  of  trees  with  an  insect  net  for  Capsids  caught  two  young 
Hyla  femorahs"  (A.  H.  Wright,  Ga.,  1931,  p.  275). 

Size:  Adults,  1-1%  inches.  Males,  24-37  mm.  Females,  23-40  mm. 

General  appearance:  This  frog  is  commonly  a  deep  reddish  brown  in  color, 
but  may  be  gray  or  greenish  gray.  It  resembles  a  small  common  tree  toad  but 
is  more  slender  and  the  black  markings  do  not  form  a  regular  cross.  There  are 
orange  or  grayish  white  spots  on  the  rear  of  the  thighs.  The  under  parts  are 
white.  The  throat  may  be  dark.  The  upper  surface  has  occasional  granula- 
tions, the  under  parts  are  areolate,  granulate  on  the  throat. 

"Cope  (1889,  p.  371)  writes:  'Body  short,  rather  broad,  and  the  entire  ap- 
pearance as  to  pattern  of  color  and  shape  not  very  dissimilar  from  Hyla  verst- 
color,  from  which,  however,  it  is  readily  distinguishable  by  the  femoral  yel- 
low spots;  the  dark  postocular  vitta,  the  absence  of  light  spots  under  the  eyes.' 

"Brimley  (1907,  p.  1*58)  characterizes  it  as  follows:  'Back  .  .  .  markings  do 
not  form  an  X-shapcd  mark.  Back  of  thigh  with  yellow  spots  or  variegations. 
No  light  spot  below  eye.  No  yellow  spots  on  sides.' 

"Deckcrt  (1915,  p.  3)  who  has  studied  southeastern  frogs  more  intimately 
than  most  observers  says  this  species  'resembles  our  own  gray  tree  frog,  with 
its  rough  skm  and  star-shaped  dark  patch  on  the  back,  but  is  smaller  and  more 
slender/ 

"We  would  consider  this  species  a  small  species.  Cope  considers  35  mm. 
above  the  average  size  and  his  largest  is  39  mm.  Le  Come  a  century  ago  gave 
1 1/£-i  %  inches  as  the  adult  range  and  it  has  been  repeated  for  this  little  under- 
stood form.  It  may  reach  45  mm.,  as  he  says,  but  of  the  140  specimens  from  20 
mm.  upwards  we  have  none  over  40  mm.,  the  average,  30  mm."  (A.  H. 
Wright,  Ga.,  1931,^273). 

"The  young  newly  transformed  frogs  arc  often  green  and  both  in  this  color 
phase  and  when  brown  arc  liable  to  be  mistaken  for  Hyla  sqtiirella  from 
which  species  they  can  always  be  distinguished  by  the  yellow  spots  on  the 
back  of  the  thighs"  (C.  S.  Brimley,  N.C.,  1940,  pp.  22-23). 

Color:  June  5,  1921.  Male.  Cephalic  half  of  pectoral  fold  pale  vinaceous- 
drab  to  darkish  grayish  brown;  posterior  half  like  the  belly.  Throat  darker 
than  the  belly. 

Female.  General  color  sorghum  brown,  deep  brownish  drab,  or  mars  brown 
on  back.  Black  spot  between  eyes.  Another  with  four  points,  two  behind  and 
two  ahead,  the  cephalic  ends  above  tympanum.  Two  on  either  side  of  the 
middle  of  the  back  and  one  over  the  crupper.  Narrow  black  or  deep  brown 
line  from  snout  through  nostril  to  eye,  and  through  tympanum  to  groin, 
where  it  breaks  up  into  spots.  Another  of  the  same  color  on  back  of  forefoot, 
forearm,  front  of  foreleg,  and  over  vent.  Below  white  line  of  the  vent  is  a 
black  one.  Spots  on  rear  of  thighs  orange  to  orange  chrome  or  light  cadmium. 
One  female  with  grayish  white  instead  of  orange  spots.  Under  parts  white 


TREE  FROGS:  HYLIDAE  321 

Pectoral  fold  pure  white  to  angle  of  the  mouth.  Chin  white  with  fine  black 
spots.  Iris  ecru-drab,  drab-gray,  or  pale  vinaceous-drab  with  reticulation  of 
black. 

Structure:  Upper  surface  with  occasional  scattered  granulations;  belly,  un- 
der surface  of  thigh,  and  breast  strongly  areolate;  throat  granulate;  pectoral 
fold  smooth  or  slightly  granulate.  Tympanic  fold  usually  present.  Muzzle 
rounded. 

Holbrook  (Gen.,  1842,  IV,  128)  alluded  to  Dumeril  and  Bibron's  mistake 
as  follows:  "Dumeril  and  Bibron  consider  this  animal  as  identical  with  the 
Hyla  sqitirella,  from  which  it  is,  however,  perfectly  distinct." 

In  1856  John  Le  Conte  (Gen.,  1856,  p.  428)  referred  to  the  same  matter.  "It 
is  wrong  in  Dumeril  and  Bibron  to  say  that  this  species  is  a  variety  of  Hyla 
squirella.  In  shape  and  size  the  difference  is  not  considerable.  The  latter  ani- 
mal during  the  warm  season  is  always  to  be  met  with  about  the  houses,  the 
Hyla  femorahs  never.  Besides,  their  notes  are  entirely  different."  As  late  as 
1882  Dumeril  and  Bibron's  mistake  somewhat  influenced  Boulenger  (Gen., 
1882,  p.  398)  in  his  statement,  "Appears  to  be  specifically  distinct  from  H.  squi- 
rella." 

Voice:  A  peculiar  cicada  note  in  chorus,  the  chorus  one  continuous  stridu- 
lating  dm  going  down  the  piny  woods  like  a  wave.  When  high  in  the  trees, 
this  frog  calls  ^<?^  at  intervals.  When  rain  brings  the  frogs  close  to  the  water 
and  the  evening  congress  is  on,  the  calls  are  speeded  up.  There  may  be  as 
many  as  6-7  a  second,  with  60-70  calls  in  rapid  succession  without  deflation  of 
the  throat. 

"Le  Conte,  1856,  seems  to  be  the  first  to  note  that  it  differed  from  H.  squi- 
rella  in  Voice.  No  one  since  has  remarked  about  its  voice  until  Deckert  (Fla., 
I9I5>  PP-  3"4)  wrote,  The  noise  resulting  from  the  calls  of  the  males  on  these 
occasions,  is  deafening.  This  call  cannot  be  reproduced  on  paper,  being  a  rapid 
succession  of  harsh,  rattling  notes  higher  in  pitch  than  the  call  of  H.  sqitirella, 
and  kept  up  all  night.  During  the  dry  season  this  tree  toad  occasionally  calls 
from  the  tops  ot  the  pine  trees,  one  answering  the  other.' 

"In  1921  the  first  general  calling  of  Hyla  femoralis  came  May  14.  Then  the 
author  went  to  Billy's  Lake  Landing  and  worked  eastward.  Heard  plenty  of 
Hyla  jemorahs  in  trees.  They  are  approaching  or  are  on  the  edge  of  cypress 
ponds.  Some  are  yet  high  in  the  trees.  They  are  calling  at  intervals  all  over  the 
piny  woods,  particularly  near  the  edges  of  cypress  bays.  One  call,  an  interval, 
one  very  faint  call,  interval. 

"On  May  16,  when  a  threatening  storm  passed  over,  Hyla  femoralis  from 
the  trees  were  almost  in  chorus.  Around  cypress  ponds  several  calling.  One 
was  on  a  projecting  piece  of  pine  bark  on  the  tree  and  within  reach,  about  six 
feet  from  the  ground.  Harper  saw  it  croak.  Afterwards  the  throat  pulsated  all 
the  time.  Harper  notes  that  it  'let  me  come  up  in  plain  sight,  within  a  couple 
of  yards  and  croaked  for  me  on  its  bark  slab-perch.  Throat  kept  distended 


322  HANDBOOK  OF  FROGS  AND  TOADS 


while  its  sides,  or  rather  whole  body,  vibrated.  Ke\9  J(e^t  about  20  times, 
usually  ending  in  %ra\,  \ra^!  Later  in  the  evening  we  visited  this  croaker  but 
it  was  gone.  Soon  we  understood.  In  two  or  three  trees  low  down  we  heard 
Hyla  femoralis.  Their  notes  are  speeded  up,  more  extended  and  continuous 
than  when  high  in  the  pines.  One  we  found  on  the  moss.  The  instant  we  en- 
tered another  pond  we  heard  a  queer  note  and  it  was  from  the  trunk  of  a  bay 
tree  right  near  us.  It  proved  to  be  a  male  Hyla  femoralis.  All  around  us  they 
were  calling.  One  was  in  a  bush  three  feet  above  ground.  Another  on  moist 
ground  at  edge  of  pond  in  amongst  six-foot  sedges.  When  a  male  croaks  it  is 
the  lower  throat  which  swells  out;  the  chin  region  does  not.  Brought  back 
three.  .  .  . 

"During  a  congress  when  several  species  are  breeding  in  the  same  pond  the 
machine-gun  calls  of  Hyla  jemoralis  make  it  difficult  to  hear  or  time  other 
calls.  Once  when  we  were  timing  the  intervals  in  Mtcrohyla's  calls,  Hyla  fe- 
moralis  calls  broke  in  frequently.  In  1921  and  1922  we  recorded  several  in- 
stances where  their  calls  drowned  out  those  of  Hyla  gratiosa,  Hyla  squirdla, 
Bitfo  quercicus,  and  Psetidacris  oculans.  We  tried  the  experiment  of  half  clos- 
ing our  ears  to  shut  out  the  Hyla  femoralts  calls.  The  other  sounds  came  out 
very  distinctly  in  the  attempt"  (A.  H.  Wright,  Ga.,  1932,  pp.  277-278). 

Breeding:  This  species  breeds  from  April  20  to  Sept.  i.  The  eggs  are  in 
groups  of  small  films  on  the  surface  or  just  below  it  attached  to  grass  blades 
of  floating  roots.  The  jelly  is  loose  and  sticky,  the  eggs  arc  brown  and  yellow- 
ish, theii  size  Ko-^n  inch  (0.8-0.9  mm.),  the  inner  envelope  Via-Viz  inch 
(1.4-2.0  mm.),  the  outer  envelope  not  distinct,  %-%  inch  (4-6  or  8  mm.). 
The  eggs  hatch  in  3  days.  The  tadpole  is  small,  i%  inches  (33  mm.),  its  tail 
tip  acuminate  and  free  of  spots,  the  lower  musculature  with  a  light  stripe. 
Many  have  bright  red  in  their  tails.  The  tooth  ridges  are  %.  After  50  to  75 
days,  the  tadpoles  transform  from  June  16  to  October,  at  }•>  inch  (13  mm.). 

"We  found  breeding  congresses  in  grassy  transient  pools  near  the  roads  in 
the  piny  woods,  in  open  ponds  in  cutovcr  roads,  in  pools  or  ditches  beside  the 
railroad  and  roads,  in  cypress  ponds  and  in  cypress  bays.  We  found  mated 
pairs  in  overflowed  grassy  fields,  shallow  transient  depressions,  in  temporary 
overflows  or  drenched  cultivated  fields,  swamps  or  dreens  in  the  cypress  bays. 
Tadpoles  were  taken  in  pools  beside  Indian  mounds,  railroads,  roads,  in  cy- 
press ponds  and  sometimes  on  the  prairies,  in  diverse  ponds  on  the  east  main- 
land or  in  bays  outside  the  swamp"  (A.  H.  Wright,  Ga.,  1932,  p.  284). 

"Dangers.  This  species  lays  in  any  pine  barrens  pool.  Many  a  shallow  grassy 
pool  will  have  packets  of  their  eggs.  Many  are  caught  by  hatching;  certainly 
many  of  the  tadpoles  never  mature.  There  must  be  a  great  loss  in  this  species. 
On  July  3,  1921,  they  were  laying  in  the  flooded  furrows  of  cane,  corn  and 
sweet  potato  fields.  There  all  were  lost.  On  June  4,  1921,  we  made  the  notes 
Tonight  the  frogs  of  several  species  (Hyla  femoralis  included)  are  laying  in 
all  sorts  of  transient  places'  "  (same,  p.  287). 


TREE  FROGS:  HYLIDAE 


3*3 


Journal  notes:  May  19,  1921,  Okefinokee  Swamp,  Ga.  On  my  way  back 
from  a  trip  to  Crosby  Pond,  at  6  P.M.  near  the  remains  of  an  old  cypress  pond 
in  piny  woods  saw  a  female  Hyla  femomlis  hopping  along  into  saw  palmetto. 
It  was  as  whitish  gray  as  any  Hyla  versicolor  I  ever  saw.  The  spot  in  middle  of 
back  showed  beautifully,  also  spot  between  the  eyes.  .  .  .  This  female  I  took 
out  to  look  at.  It  leaped  away  onto  the  gray  sand.  Had  a  hard  time  seeing  it  be- 
cause it  matched  the  gray  sand  so  well.  In  one  minute  since  its  capture,  it  had 
darkened  considerably. 

In  denser  cover  9  inches  high  with  small  saw  palmetto  and  small  bushes 
found  a  half-grown  Hyla  femorahs.  It  was  green  on  its  back  (very  suggestive 
of  H.  sqinretta,  which  strangely  enough  we  don't  get  here). 

May  21,  in  the  compartment  of  Hyla  femoralis,  most  of  the  specimens  that 
were  green  when  captured,  in  fact  all  (including  one  little  half-grown  one), 
are  now  Vandyke  brown  or  moss  brown.  Adults  are  not  often  green,  but  their 
life  at  transformation  frequently  starts  in  a  green  livery. 

A  captive  female  in  a  jar,  June  19,  is  pale  light  mouse  gray  on  back  with  no 
markings. 

On  April  23,  1921,  the  boys  found  two  on  a  rail  fence  at  2:30  P.M.  The  next 
day  they  brought  three  more  from  the  same  fence.  On  April  26,  the  boys 
found  some  more  Hyla  femorahs  m  the  rain  barrels  along  the  railroad  and 
near  the  lumber  company's  woodpile.  In  a  pine  near  camp  about  15  or  20  feet 
up  on  the  end  fork  of  a  large  branch  is  a  Hyla  femoralts  male.  It  doubtless  is 
the  one  we  have  heard  ever  since  we  have  been  here. 

May  1 8,  their  calls  made  a  perfect  dm  in  cypress  bays  and  north  edge  of 
Long  Pond.  Some  were  m  bushes,  others  in  bay  trunks,  and  others  were  hop- 
ping in  moss  edges  or  grassy  edges  of  the  pond  or  among  the  lizard's-tail. 
Another  congress  June  4,  1921.  On  one  palmetto  within  2  or  3  feet  of  each 
other  were  five  male  H.  fenwralis  calling. 

July  11,  1922,  7:30-11  P.M.  We  heard  no  end  of  Hyla  femoralts  in  every 
crossing  and  cypress  pond.  .  .  .  They  were  especially  centered  about  a  clump 
of  saw  palmettos.  Here  Miles  Pirnie  tound  one  pair  and  I  another.  Later, 
on  one  blade  were  two  more  pairs  and  one  male.  In  this  clump  were  possibly 
20-^0  males  calling.  This  breeding  came  after  a  hard  rain.  The  females  doubt- 
less do  not  enter  the  ponds  until  about  ready  to  lay. 

July  6.  Mrs.  Chesser  hoed  out  another  female  in  the  open  field.  There  has 
been  a  chorus  of  them  m  the  pond  near  by.  Probably  the  female  was  resting 
here  during  the  day. 

Authorities'  corner: 
J.  E.  Holbrook,  Gen.,  1842,  IV,  128. 
R.  Dcckert,  Fla.,  1915,  p.  3. 
A.  H.  Wright,  Ga.,  1932,  pp,  293-294. 

One  of  the  least  known  forms  of  the  Coastal  Plain  is  Hyla  femoralis,  which 
we  have  always  associated  with  cypress  ponds,  bays,  and  thickets.  A  very  sig- 


324  HANDBOOK  OF  PROGS  AND  TOADS 

nificant  extension  of  its  range  comes  in  its  recorded  occurrence  by  J.  A.  Fowler 
and  Grace  Orton  (Md.,  1947,  p.  6)  at  Battle  Creek,  Maryland. 

Barker,  Barking  Frog,  Coat  Bet,  Florida  Tree  Frog,  Georgia  Tree  Frog, 
Florida  Hyla,  Bell  Frog,  Giant  Tree  Frog 

Hyla  gratiosa  Le  Conte.  Plate  LXIX;  Map  19. 

Range:  North  Carolina  (B.  B.  Brandt)  to  Florida  to  Louisiana.  Hartselle, 
northern  Alahama  (Cahn). 

Habitat:  Trees  of  hammocks,  pine  barrens,  and  bays.  Breeds  in  pine  bar- 
ren ponds  and  cypress  ponds. 

"High  pine,  high  hammock,  and  dry  flatwoods,  in  the  upper  branches  of 
long-leaf  and  slash  pine  and  of  live-oak.  .  .  .  Uncommon  locally  and  gener- 
ally. Wright  (1953)  describes  the  rain-song;  this  odd  call  may  be  heard  rarely 
from  the  treetops  in  high  pine.  On  April  u,  1933,  H.  K.  Wallace  and  I  dug 
two  adults  and  a  yearling  out  of  slightly  moist  sand  more  than  four  feet  deep 
in  an  Indian  mound,  in  cutover  pmeland  in  Lake  County.  On  November  24, 
1935,  I  found  another  in  the  same  mound  at  about  the  same  depth.  This 
winter-burrowing  may  be  of  significance  in  explaining  the  fact  that  gratiosa 
is  almost  never  seen  except  during  warm  weather"  (A.  F.  Carr,  Jr.,  Fla.,  1940^ 
p.  60). 

Size:  Adults,  2-2%  inches.  Males,  49-68  mm.  Females,  50-68  mm. 

General  appearance:  This  is  our  largest  native  tree  frog.  It  is  ashen  gray, 
purplish,  or  green  in  color.  The  skin  is  evenly  granular,  the  back  evenly  cov- 
ered with  elliptical  or  round  spots  darker  than  the  general  color  and  encircled 
with  black.  These  spots  may  be  absent  in  some  of  the  color  phases  this  frog 
assumes.  A  light  stripe  extends  along  the  sides,  bordered  below  by  a  purplish 
brown  one.  There  is  some  yellow  on  the  sides  in  axilla  of  arm  and  in  the  groin. 
The  under  parts  are  creamy  or  pinkish  white.  The  throat  of  the  male  is  green 
or  yellow  with  dark  spots  just  back  of  the  chin,  while  that  of  the  female  has 
the  center  throat  light,  and  the  sides  of  throat  and  pectoral  region  sulphur 
yellow.  The  colored  area  of  throat  is  encircled  with  white  on  the  inner  side. 
The  spots  on  the  side,  chin,  and  rim  of  jaw  are  reddish  brown. 

Color:  "Mule.  June  5,  1921.  Lettuce  green  on  hind  legs  and  fore  legs,  oil 
green  or  cerro  green  on  the  back.  The  encircled  spots  of  the  back  have  a  ring 
of  mars  brown  or  bone  brown  or  other  browns  with  green  ground  color 
within.  There  are  occasional  spots  as  big  as  pin  heads  or  twice  so  of  bright 
green-yellow  or  greenish  yellow  or  green-yellow.  This  color  is  in  the  groin 
extending  to  just  back  of  arm  insertion  where  for  one  half  an  inch  is  a  clear 
whitish  area.  The  greenish  yellow  from  groin  forward  demarcates  side  color 
from  pinkish  white  or  creamy  belly  color.  From  tip  of  snout  along  upper  jaw 
under  tympanum  and  along  side  to  within  i  inch  or  %  inch  of  leg  insertion 
is  a  pinkish  white  stripe.  Below  it  is  a  line  of  mars  violet  or  taupe  brown  (pur- 


TREE  FROGS:  HYLIDAE 


325 


Plate  LXV11I.  Hyla  femoralis.  1,2.  Males 
calling  (X'/$).  3-  Tadpole  (Xi%)-  4-  Egg 
cluster  (XO.  5-  Male  (Xi%)-  6.  Female 
(Xi).  7-  Female  (Xi%). 


LX/X.  Hyla  gratiosa,  1,7.  Males 
(X%).  2.  Male  croaking  (X#).  3-  Male 
in  a  cornfield  at  night  (X1^)-  4-  Female 
(X#).  5-  Transforming  frog  (X%)«  6. 
Eggs  (X%). 


326  HANDBOOK  OF  FROGS  AND  TOADS 

plish  brown)  which  expands  behind  the  angle  of  the  mouth  into  a  large  area 
on  the  side.  Same  color  back  of  pinkish  white  or  white  area  back  of  brachium 
and  antebrachium  onto  the  last  finger.  Same  white  line  around  vent  with 
mars  violet  below.  Same  combination  on  the  knee.  Two  or  three  white  patches 
on  foot  with  mars  violet  between  and  behind  them.  Back  of  femur  or  thigh 
dull  Indian  purple.  No  spots.  Below  the  expanded  mars  violet  area  of  the  side 
is  a  small  whitish  line  %-i  inch  long,  then  comes  greenish  yellow  and  finally 
the  belly  color.  Just  back  of  the  chin  is  grayish  white  speckled  with  taupe 
brown;  then  lettuce  green  or  oil  green  from  angle  of  mouth  around  to  angle 
of  mouth,  the  band  Va  inch  wide  when  not  inflated;  the  wrinkled  part  light 
orange  yellow  or  deep  chrome.  This  color  extends  to  pectoral  fold  of  skin  from 
forearm  to  forearm.  Ins  spotted  purplish  black  and  bronzy  or  vinaceous. 

"Males,  ...  in  botany  drum.  Upon  opening  the  can  one  was  all  green 
with  a  few  yellowish  spots  but  not  the  regular  circles,  the  stripe  along  the 
jaw  and  on  the  side  reminding  one  of  the  same  in  Hyla  ctnerea.  Hardly  any 
white  or  purplish  shows  anywhere.  Another  was  a  vinaceous  gray  with  the 
rings.  Another  dark  green  and  the  others  still  different. 

"Female.  Belly  and  under  side  of  legs  white,  a  few  picric  yellow  spots  on 
side  below  band,  also  rear  end  of  white  line  on  side  becomes  picric  yellow  in 
the  groin.  Picric  yellow  in  the  axilla  and  on  cither  side  of  light  central  throat 
and  chin  region  which  is  white  or  sulphur  yellow  with  a  few  spots.  Pectoral 
region  sulphur  yellow.  Stripe  around  snout  along  upper  jaw  to  groin  white 
except  for  rear  and  not  interrupted  as  in  the  male.  Spots  on  back  black  encir- 
cled. Color  of  side  and  chin,  spots,  rim  of  jaw,  bands  on  arms  hays  maroon, 
chocolate,  warm  sepia  or  bone  brown.  Upper  parts  calliste  green,  cosse  green 
or  apple  green"  (A.  H.  Wright,  Ga.,  19^2,  p.  302). 

Structure:  A  large  tree  frog  of  heavy  build;  head  broad  and  short;  fingers 
webbed;  large  disks  on  fingers  and  toes;  skin  has  marked  secretion  giving 
strong  persistent  odor.  Upper  parts  except  arms  and  legs  strongly  granulate; 
under  surface  of  hind  limbs  granulate  only  on  posterior  half  of  thigh,  other- 
wise smooth;  under  surface  of  arm,  prominent  breast  fold,  and  lower  throat 
smooth  or  slightly  granulate;  chin  granulate;  webbing  of  hind  feet  extends  to 
the  disks  which  are  very  large,  3  mm.,  or  0/5  of  tympanum;  disks  of  fore- 
feet, 4  mm.,  or  0.66  of  the  tympanum;  a  dermal  fold  over  the  ear  to  the  shoul- 
der and  continued  often  as  a  loose  fold  along  the  side  of  the  body  halfway 
to  the  groin;  a  tarsal  fold;  vomerine  teeth  between  inner  nares;  head  ahead 
of  eye  short,  obtuse;  vocal  sac  well  developed. 

"During  the  last  spring,  whilst  I  was  residing  in  the  lower  country  of 
Georgia,  it  was  my  good  fortune  to  meet  with  three  specimens  of  the  animal 
described  below.  One  of  them  was  taken  in  the  water  of  a  pine  barren  pond, 
another  was  found  in  a  cavity  of  a  sand  pit,  and  the  third  upon  a  tree  in  the 
forest. 

"This  Hyla  is  remarkable  for  its  size,  approaching  in  this  respect  to  those 


TREE  FROGS:  HYLIDAE  317 

found  in  tropical  regions.  Two  of  them  were  of  a  greenish  dusky;  the  second, 
who  had  concealed  himself  in  a  hole  in  the  sand,  was  of  a  bright  pea  green, 
but  in  the  space  of  half  an  hour  he  changed  to  the  color  of  the  others,  thus 
showing  a  complete  possession  of  the  faculty  of  changing  color  at  will,  so 
remarkable  in  many  of  the  Batrachia. 

"There  yet  remains  undiscovered  and  undescnbed  in  Georgia  three  species 
of  this  genus,  which  have  as  yet  eluded  my  search.  The  notes  of  these  are  re- 
markably distinct  from  those  of  others;  I  may  hereafter  be  fortunate  enough 
to  obtain  them. 

"Coarsely  granulate  both  above  and  beneath.  Color  above  varying  at  the 
will  of  the  animal  from  bright  green  to  cinereous  and  to  greenish  dusky,  with 
roundish  spots  or  irregular  blotches  of  darker,  or  speckled  with  variously 
shaped  dots  of  the  same,  all  of  them  with  some  few  small  yellow  irregularly 
disposed  spots  on  the  back  and  sides.  Beneath  whitish,  more  or  less  inclining 
to  yellow  or  orange.  Upper  lip  white,  or  white  varied  with  green  or  dusky; 
lower  lips  sometimes  whitish,  in  others  of  the  color  of  the  back;  in  some  a 
white  line  extends  from  the  upper  lip  along  the  side  to  the  insertion  of  the 
hind  leg,  in  others  the  sides  are  more  with  rounded  spots  of  darker  and  no  line 
visible.  Irides  black  varied  with  golden;  tympanum  copper-colored,  a  consid- 
erable depression  between  the  nostrils  and  the  eyes.  Chin  varied  with  dusky 
or  green,  with  a  slight  fold  at  the  bottom;  transverse  space  between  the  arms 
smooth,  without  any  granulations.  Arms  and  legs  barred,  with  darker,  yellow- 
ish or  reddish  on  the  under  side,  the  former  smooth  beneath,  the  latter  granu- 
late on  the  posterior  half;  the  under  side  of  the  posterior  half  of  the  thighs  is 
smooth.  Disks  of  the  toes  very  large. 

"Length  of  head  and  body  2.5  inches;  humerus  .6;  antcbrachium  .6;  hand 
.75;  femur  1.2;  tibia  1.15;  foot  1.6"  (J.  Le  Conte,  Ga.,  1856,  p.  146). 

Voice:  The  call  is  woody,  deep,  a  curious  tonl^f  ton^,  like  someone  pounding 
on  a  hollow,  heavy  barrel  or  hogshead.  The  call  in  the  ponds  is  coat  bet.  The 
call  from  trees  as  it  approaches  water  is  a  bark.  The  vocal  sac  is  a  large  sub- 
gular  vocal  pouch. 

The  first  time  we  heard  this  species  to  know  it  as  such  was  on  June  5, 1921. 
After  a  period  of  heavy  rainfall,  we  visited  in  the  evening  an  oak  toad  pool 
near  the  cultivated  fields  of  Billy's  Island,  Ga.,  and  heard  this  curious  call.  It 
seemed  at  first  somewhat  like  the  call  of  Rana  clamitans  only  too  many  times 
separated,  and  actually  the  resemblance  is  fleeting  and  slight.  There  are  55-60 
calls  a  minute,  the  throat  often  remaining  partly  inflated  between  calls. 

Breeding:  This  species  breeds  from  March  to  August.  The  eggs  are  laid 
singly  on  the  bottom  of  the  pond.  The  single  envelope,  Mo-%  inch  (2.3-5.0 
mm.),  is  loose,  glutinous,  and  indefinite  in  outline.  The  vitelline  membrane 
appears  as  an  inner  envelope,  Kc-Ko  inch  (1.6-2.5  nun.)  in  diameter,  the 
egg  %5-Mfl  inch  (1.0-1.8  mm.).  The  greenish  tadpole  is  medium,  2  inches 
(50  mm.)  long,  and  is  the  largest  Hylid  tadpole  of  the  eastern  United  States. 


328  HANDBOOK  OF  FROGS  AND  TOADS 

The  tail  is  long,  its  tip  acuminate  with  a  flagellum,  and  the  tooth  ridges  are 
%.  The  tadpole  period  is  40  to  70  days.  The  tadpoles  transform  from  July  to 
October,  at  %G-%  inch  (14-20  mm.). 

"Choruses  of  more  than  twenty  or  twenty-five  males  are  not  often  seen. 
They  call  from  deep  water  in  small  permanent  ponds,  from  March  3  to  August 
14.  In  Gainesville  the  first  choruses  have  always  gathered  in  the  power  plant 
water  aerator,  where  the  water  is  abnormally  warm;  here  I  have  also  seen 
them  feeding  on  insects  at  the  lighted  entrance"  (A.  F.  Carr,  Jr.,  Fla.,  1940^ 
p.  61). 

Journal  notes:  While  we  were  in  the  Okefinokee  Swamp,  Ga.,  we  usually 
recorded  this  species  as  Barker,  Barking  Frog,  or  Coat  Bets.  The  last  refers  to 
the  normal  note  in  the  breeding  pools,  the  others  to  a  puzzle  that  perplexed  us 
for  two  seasons. 

On  July  15,  1921,  on  Chesser  Island  we  heard  of  Coat  Bet  frogs,  so  named 
from  the  sound  of  their  breeding  call.  On  July  16,  during  the  morning,  we 
heard  a  barking  frog  in  the  trees  south  of  camp.  That  night  in  a  near-by  pond 
was  an  immense  chorus  of  Coat  Bets.  In  1922  the  barkers  perplexed  us  even 
more. 

Not  until  July  26  did  we  solve  the  puzzle.  About  three  miles  along  the  road 
from  Chesser  Island  to  Folkston  we  heard  in  the  evening  in  a  cypress  pond 
to  the  right  of  the  road  some  Hyla  gratwsa,  and  beyond  them,  a  barker  or 
two.  We  went  after  the  barker,  and  found  one  in  a  small  gum  4-5  feet,  pos- 
sibly 6  feet  up.  It  is  Hyla  gratwsa!  I  saw  him  do  it.  There  were  two  more 
barkers  besides  the  one  I  caught.  Several  Hyla  gratiosa  were  in  the  water 
calling  normally.  Coat  Bets  and  barkers  are  one. 

June  5,  1921,  Okefinokee  Swamp.  Later  we  found  other  croakers.  One  was 
on  the  raised  ndgc  of  land  beside  a  cornstalk.  It  was  green  with  the  spots 
usually  figured.  Another  male  was  lying  flat  in  the  water  at  the  edge  of  a 
weedy  ridge.  It  was  absolutely  flat  and  spread  out.  It  was  without  the  spots 
and  uniform  dull  brownish  green,  like  the  color  of  the  water.  Another  was  in 
a  furrow  between  two  rows  of  corn.  It  was  spotted  and  alert.  The  first  one 
rested  on  ground  more  or  less  horizontally;  the  second  in  water  horizontally. 
And  the  third  more  or  less  diagonally  upright.  The  call  can  be  heard  at  a  long 
distance.  Usually  the  heels  are  widely  separated  at  croaking  and  the  lower  part 
of  the  body  dips  somewhat.  Found  a  male  Hyla  gratiosa  beside  a  grassy  bank. 
He  was  as  big  and  round  as  the  can  top  of  a  mason  jar.  How  his  throat  would 
pu(T  out  I 

Authorities9  corner: 
R.  F.  Deckert,  Fla.,  1915,  pp.  4-5. 
C.  J.  Coin,  Fla.,  1938,  p.  48. 
A.  R.  Cahn,  Ala.,  1939,  pp.  52-53. 


TREE  FROGS:  HYLIDAE  329 

Pacific  Tree  Frog,  Pacific  Tree  Toad,  Pacific  Hyla,  Wood  Frog  (Cooper), 

Pacific  Coast  Tree  Toad 

Hyla  regilla  Baird  and  Girard.  Plate  LXX;  Map  18. 

Range:  Vancouver  Island  and  British  Columbia  to  Lower  California;  east 
to  western  Montana  (Rodgcrs  and  Jellison),  Idaho,  Utah,  Nevada,  and  Flag- 
staff, Ariz. 

Habitat:  On  the  ground,  especially  about  streams,  springs,  ponds,  swamps, 
and  other  moist  places;  irrigation  ditches. 

"Extends  to  timber-line  in  the  Sierra  Nevada;  occurs  in  all  zones  below 
Alpine-Arctic.  Inhabits  damp  recesses  among  rocks  and  logs;  the  ground  in 
the  vicinity  of  springs,  streams  and  lakes;  rank  growth  of  vegetation,  espe- 
cially in  marshy  places,  trees  in  damp  forests;  and  in  open  country,  burrows  of 
various  animals"  (J.  Grinnell  and  C.  L.  Camp,  Calif.,  1917,  p.  145). 

"This  tree-toad  is  probably  the  most  abundant  batrachian  in  California, 
where  it  ranges  from  sea-level  to  n,6oo  feet.  It  may  be  found  abundantly  in 
marsh-lands,  lakes,  springs,  under  the  bark  of  trees  and  in  almost  any  other 
place  where  there  is  continued  moisture.  It  does  not  appear  to  congregate  in 
such  large  compact  masses  as  does  Hyla  arenicolor,  but  may  be  fo.und  in  large 
colonies  during  the  breeding  season"  (J.  R.  Slevin,  B.C.,  1928,  p.  115). 

"Rio  Santo  Domingo  at  the  Hamilton  Ranch,  300  ft.,  lat.  ^o°45/;  37249- 
37250;  June  2.  Aguaito  Springs,  15  miles  E.  Rosario,  1300  ft.,  lat.  30°4';  37251- 
37256;  June  9.  On  the  night  of  June  2  multitudes  of  Pacific  tree  toads  were 
strung  along  the  sand  bars  and  along  the  sandy  banks  of  the  shallow,  widely 
spread,  slow-moving  Santo  Domingo  River.  Their  croaking  was  the  domi- 
nant night  sound.  When  the  toads  were  alarmed,  they  would  swim  to  and 
try  to  hide  in  the  masses  of  slimy,  thread-like  green  algae  attached  to  fallen 
willow  branches  and  other  debris.  The  species  was  also  abundant  at  night  in 
and  around  the  isolated  pools  at  Aguaito  Springs.  Individuals  beside  but  not 
actually  in  the  water,  if  disturbed,  hopped  uphill  to  dry  brush  cover  rather 
than  down  into  the  water"  (L.  Tevis,  Jr.,  L.C.,  1944,  P-  ?)• 

Size:  Adults,  T-I%  inches.  Males,  25.5-48.0  mm.  Females,  25-47  mm-  ^n 
the  whole,  the  mature  females  are  larger  than  the  males,  except  occasional  un- 
usually large  males. 

General  appearance:  This  small,  delicate  tree  toad  is  somewhat  smaller  and 
more  slender  than  the  canyon  tree  toad  (Hyla  arenicolor)^  which  like  this 
form  has  the  rear  of  the  thighs  uniform,  not  spotted.  This  species  reminds 
the  authors  of  species  of  Pseudacris.  The  disks  on  the  fingers  and  toes  are 
larger,  however.  It  is  very  variable  in  color,  usually  with  stripes  on  the  back 
and  a  triangle  between  the  eyes,  and  also  with  a  stripe  along  the  side  of  the 
head. 

One  male  is  light  brown  in  color  with  a  dark  brown  V  between  the  eyes 
and  two  rows  of  large  dark  spots  on  the  back.  It  has  a  conspicuous  greenish 


330  HANDBOOK  OF  FROGS  AND  TOADS 

black  line  from  the  nostril  to  the  eye  and  from  the  eye  through  and  beyond 
the  tympanum.  Then  there  is  a  broken  line  of  spots  to  the  groin.  There  is 
orange  or  yellow  in  the  groin  and  on  the  rear  of  femur  and  on  the  foot.  The 
arms,  legs,  and  feet  are  indistinctly  barred  with  dark.  The  upper  jaw  is  a 
beautiful  light  pinkish  cream  color.  The  throat  of  this  male  is  dark  in  color, 
greenish.  In  the  middle  of  the  light  belly  is  a  broad,  longitudinal  bluish  area. 

Another  male  is  bright  green,  with  an  indistinct  triangle  between  the  eyes 
and  round  dark  spots  on  the  back.  The  dark  mask  is  bordered  above  by  a 
light  pinkish  cream  line  and  below  by  the  light  jaw.  The  throat  is  olive,  with 
some  orange  on  the  center  rear  portion.  Orange-yellow  is  conspicuous  in  the 
groin  and  on  the  rear  of  the  femur. 

"None  of  the  thirty-one  specimens  from  the  vicinity  of  Lake  Como  has  a 
color  pattern  that  approaches  the  uniform  bold  pattern  of  the  Hyla  regilla 
from  the  Great  Basin.  Most  of  them  more  nearly  resemble  specimens  from 
Washington,  Oregon  and  California.  This  evidence  tends  to  confirm  what 
might  be  expected,  namely,  that  the  population  of  the  Bitterroot  Valley  is 
related  to  that  of  the  Pacific  Coast  through  past  or  present  connection  along 
the  Columbia  River  drainage.  Fifteen  of  the  specimens  have  a  color  pattern 
that  seems  to  be  peculiar  to  the  Bitterroot  Valley  in  that  it  is  broken  up  into 
many  small  units.  One  specimen,  the  most  extreme  in  this  respect,  has  88  small 
dark  gray  spots  of  irregular  shape  on  the  back.  The  heavy  Y-shaped  mark  be- 
tween the  eyes  that  is  so  characteristic  of  Hyla  regtlla  is  represented  on  some 
of  these  specimens  only  by  a  narrow  branched  line.  The  markings  on  the  legs 
are  also  broken  into  many  more  small  units  than  is  typical  for  the  species. 
Since  there  were  more  than  a  thousand  specimens  in  the  collection  from  the 
Pacific  Coast  area  with  which  our  specimens  were  compared,  it  seems  reason- 
able to  assume  that  all  of  the  major  color  patterns  that  might  appear  in  indi- 
viduals from  the  Pacific  Coast  were  represented,  and  from  this  it  might  fol- 
low that  the  population  around  Lake  Como  is  unique  in  having  this  finely 
speckled  color  phase"  (T.  L.  Rodgcrs  and  W.  J.  Jellison,  Mont.,  1942,  pp. 
10-11). 

Color:  Male.  Pullman,  Wash.,  from  Dana  J.  Leffingwell,  March  28,  1928. 
Upper  parts  some  variation  of  dark  grayish  olive,  olive-buff,  Isabella  color, 
old  gold,  chamois,  sulphine  yellow,  oil  yellow,  oil  green,  green,  cinnamon- 
buff,  or  clay  color.  Sometimes  with  brown  band  through  the  eye  alone,  usually 
with  bandhke  spots  of  Saccardo's  umber,  sepia,  or  brownish  olive  on  the  back 
forming  a  median  stripe  and  one  on  either  side,  also  with  triangle  between 
the  eyes,  and  also  stripe  in  front  of  eye.  Some  of  small  spots  on  sides  are  at 
times  almost  black.  Stripe  from  below  nostril  along  upper  lip  to  shoulder  in- 
sertion is  primrose  yellow.  Outside  the  long  dorsolateral  spot,  back  is  honey 
yellow  becoming  cream-buflf  or  cartridge  buff  on  sides.  Upper  parts  of  fore 
limbs  chamois  or  old  gold,  with  light  brownish  olive  crossbars.  Underside  of 


TREE  FROGS:  HYLIDAE 


33' 


arms  olive-ocher,  dark  olive-buff.  A  buff-olive  line  along  outer  edge  of  fore- 
arm. Groin  aniline  yellow  as  also  on  front  and  underside  of  femur  and  to 
some  extent  on  underside  of  tibia  and  hind  foot.  Lower  part  of  rear  of  femur 
aniline  yellow  with  upper  edge  of  medal  bronze.  There  is  also  a  dark  edge  to 
front  of  tibia.  Belly  either  side  of  meson  vmaceous-cmnamon  or  pinkish  buff; 
wash  of  same  on  underside  of  fore  and  hind  limbs.  Vocal  sac  when  inflated  is 
old  gold  or  ohve-ocher;  deflated  it  is  buffy  olive  or  olive  in  front  and  olive- 
ocher  behind.  Iris:  pupil  rim  behind  and  in  front  broken  by  black;  above  the 
rim,  lemon  yellow;  rest  of  iris,  raw  sienna  or  antique  brown. 

Female.  San  Diego,  Calif.,  from  L.  M.  Klauber,  March  28,  1928.  May  be 
like  male,  but  with  little  yellow  on  the  groin.  Front  of  femur  and  underside 
of  tibia  olive-ocher  or  aniline  yellow.  Entire  under  parts  including  throat 
may  be  cartridge  buff,  or  throat  may  be  light  with  indistinct  dots  of  olive. 
Under  parts  of  hind  legs  avellaneous  or  wood  brown  like  males. 

Another  specimen.  Las  Vegas,  Nev.,  Aug.  22,  1925.  One  half-grown  frog 
was  light  paris  green  on  upper  parts;  no  dark  spots  except  vitta;  a  little  of  clay 
color  or  cinnamon  buff  in  middle  of  back;  the  fore  and  hind  legs  mainly  light 
pinkish  cinnamon. 

Structure:  Upper  parts  smooth,  not  warty  as  is  the  usual  condition  of 
H.  arenicolor;  prominent  breast  fold;  tympanum  round. 

Original  descriptions  under  different  names:  "Hyla  regtlla,  B.  &  G.  This 
is  a  species  of  medium  size;  the  largest  individual  observed  measuring  one 
inch  and  a  half  from  the  nose  to  the  posterior  extremity  of  the  body,  the  head 
itself  occupying  about  half  of  this  length.  The  hind  legs  are  long  and  slender, 
the  web  extending  only  to  half  the  length  of  the  longest  toe;  fingers  compara- 
tively long.  The  general  color  is  green  above,  turning  to  orange  yellow  along 
the  sides  of  the  head,  abdomen  and  legs.  Two  oblong,  brownish  black  spots 
exist  on  the  occiput,  from  which  two  vittae  (one  pair)  of  the  same  black  color 
extend  along  the  dorsal  region;  a  similar  band  passes  from  the  tip  of  the  nose, 
across  the  eye  and  tympanum,  and  along  the  abdomen,  when  it  is  interrupted 
and  forms  a  series  ot  black  and  irregular  small  spots.  In  the  immature  state, 
green  is  the  prevailing  color;  a  few  black  spots  being  present  along  the  whitish 
abdomen.  Specimens  of  this  species  were  collected  on  Sacramento  River,  in 
Oregon  and  Puget  Sound.  Drawings  from  life  were  made  on  the  spot  by 
Mr.  Drayton"  (S.  F.  Baird  and  C.  Girard,  Ore.,  1854,  p.  174). 

"Litoria  occidental}*,  B.  and  G. — Throat  smooth.  Abdomen,  sides  of  body 
and  lower  surface  of  thighs  granulated.  Tympanum  very  small.  Fingers  al- 
most or  entirely  free;  toes  slightly  webbed  at  the  base;  extremities  of  both  not 
dilated.  Color  above  pale  chestnut,  with  obscure  or  obsolete  blotches  of  darker. 
Beneath  white.  A  few  crossbands  on  the  outside  of  the  legs.  A  dark  chestnut 
line  beginning  at  the  nostril,  passes  back  through  the  eye,  behind  which  it 
widens  so  as  to  include  the  tympanum,  stopping  just  above  the  insertion  of 


332  HANDBOOK  OF  FROGS  AND  TOADS 

the  arm.  One  or  two  oblique  blotches  of  dark  chestnut  on  each  side.  Body 
i Vic  inches  long;  hind  leg  extended  iVo  inch.  Hab.  San  Francisco"  (S.  F. 
Baird  and  C.  Girard,  Calif.,  1853,  p.  301). 

"Hyla  regtlla,  B.  and  G.3  Proc.  Acad.  Nat.  Sc.,  Philad.,  VI.  1852,  174.  Syn. 
Hyla  scaptdaris,  Hallow.,  Proc.  Acad.  Nat.  Sc.  Philad.,  VI.  1852,  183"  (S.  F. 
Baird  and  C.  Girard,  Calif.,  1853,  p.  301). 

"Hyla  scapularis.  Sp.  char.  Head  small;  body  small  and  slender,  olive  green 
above,  with  numerous  bluish  blotches;  a  bluish  vitta  running  from  the  eye 
over  the  shoulder;  total  length  one  and  a  half  inches  (Fr.).  Description.  The 
head  is  short  and  small,  depressed;  the  snout  somewhat  rounded;  the  nostrils 
are  small  and  circular,  looking  upwaid  and  outward,  about  a  line  apart,  situ- 
ated immediately  below  the  ridge  running  from  the  extremity  of  the  snout 
to  the  anterior  canthus  of  the  eye;  they  are  nearer  the  extremity  of  the  snout 
than  the  eye;  mouth  quite  large;  the  tongue  is  heart-shaped,  quite  free  behind, 
notched  upon  its  posterior  border;  there  are  two  series  of  palatine  teeth  be- 
tween the  nostrils,  and  separated  from  each  other  by  a  narrow  intermediate 
space;  the  eyes  are  round  and  project  considerably;  the  tympanum  is  small 
and  circular;  the  body  is  flattened,  rather  slender,  much  contracted  posteriorly; 
extremities  slender;  the  upper  surface  of  the  body  and  extremities  present 
numerous  small  granulations;  abdomen  and  under  surface  of  extremities  much 
granulated;  the  granulations  upon  the  abdomen  vary  in  size,  and  are  closely 
in  juxtaposition;  chin  and  throat  granulated.  Color.  Ground  color  above 
greenish  olive,  presenting  numerous  irregular  bluish  blotches  upon  the  sur- 
face; several  deeper  colored  blotches  upon  the  sides;  a  bluish  vitta,  about  two- 
thirds  of  a  line  in  breadth,  extends  from  the  posterior  part  of  the  eye  along  the 
sides  of  the  neck  over  the  shoulder,  a  short  distance  beyond  which  it  termi- 
nates; upper  surface  of  extremities  marked  with  bluish  spots.  Dimensions. 
Length  of  head  5  lines;  greatest  breadth  5  lines;  length  of  body  i  inch;  length 
of  humerus  4  lines;  of  forearm  3%  lines;  of  hand  to  extremity  of  longest  finger 
5  lines;  length  of  thigh  7  lines;  of  leg  8  lines;  of  foot  to  extremity  of  longest 
toe  7%  lines;  total  length  i  inch  5  lines.  Habitat.  Oregon  Territory.  Presented 
to  the  Academy  by  Dr.  Shumard"  (E.  Hallowell,  Ore.,  1854,  p.  183). 

Voice:  The  call  is  fy-ec^-e^  in  rapid  sequence. 

"This  is  the  common  frog  of  the  coastal  areas,  whose  loud  croaking  is  to  be 
heard  nightly  in  the  spring  about  every  puddle,  pond,  and  creek.  The  noise 
made  is  surprising  for  so  small  a  creature.  It  has  the  capacity  to  change  its 
color  considerably  and  may  be  bright  green,  gray  or  brown  in  almost  any 
shade"  (L.  M.  Klauber,  Calif.,  1934,  P-  ?)• 

"In  the  spawning  season,  or  more  properly,  as  long  as  there  is  any  likelihood 
of  spawning,  the  'chorus  note,'  the  note  which  serves  to  bring  the  scattered 
individuals  together  for  breeding  purposes,  is  uttered.  This,  to  human  ears, 
sounds  like  kreck-ek  and  is  uttered  over  and  over  again  in  rapid  sequence. 
...  At  any  season  of  the  year  this  hyla  may  be  heard  to  utter  a  single  pro- 


TREE  FROGS:  HYLIDAE 


333 


longed  note,  kr-r-r-eck,  lower  in  pitch  than  the  song  note.  This  note  may  be 
used,  as  indicated  above,  at  the  ending  of  a  'chorus';  but  it  is  also  uttered  in 
other  seasons  of  the  year  when  the  individuals  are  hidden  in  their  daytime 
retreats,  and  when  no  'songs'  are  to  be  heard"  (T.  I.  Storer,  Calif.,  1925,  pp. 
221-222). 

Breeding:  This  frog  breeds  from  early  January  to  mid-May.  (See  "Journal 
Notes"  for  1942.)  The  brown  and  yellowish  eggs,  in  small,  loose,  irregular 
masses  (10-70  eggs)  are  laid  in  quiet  water  beneath  or  sometimes  at  the  sur- 
face, attached  to  vegetation.  They  remind  one  of  Pseudacris  masses.  The  egg 
is  %o  inch  (1.3  mm.),  the  inner  envelope  (firm)  VJo  inch  (2  mm.),  the  outer 
envelope  (loose,  sticky)  %<\-%  inch  (4.7-6.7  mm.).  The  tadpole  is  medium, 
i%  inches  (46.6  mm.),  full  and  deep-bodied,  its  tail  tip  acute  or  obtuse  with- 
out a  flagellum.  It  is  dark  brownish  in  color  and  the  tooth  ridges  are  %.  After 
50  to  80  days  the  tadpoles  transform  from  May  15  to  September  i,  at  ^o-1/yie 
inch  (11-17  mm')'  Mrs.  Katherine  Van  Winkle  Palmer  took  ten  transform- 
ing Hyla  regilla  at  Oakvillc,  Washington,  July,  1923.  They  ranged  from  12 
to  15  mm.  Our  series  of  36  from  Alta  Meadows,  Sequoia  National  Park,  Calif,, 
Aug.  22-27,  1917,  ranged  from  11.5-17.0  mm.;  mode  13.5,  average  14.5  mm. 

"The  egg  clusters  of  this  hvla  contain  a  varying  number  of  eggs.  In  labora- 
tory aquaria  and  occasionally  in  the  field  single  eggs  are  laid.  Some  masses 
total  over  60  eggs.  Probably  the  number  in  a  mass  reflects  in  a  general  way 
the  degree  of  solitude  experienced  by  a  pair  when  eggs  are  deposited.  Where 
only  one  female  is  laying  a  few  large  clumps  are  to  be  found.  Where  many 
hylas  have  been  spawning  in  one  place  the  egg  masses  are  small. 

"At  times  very  large  numbers  of  eggs  are  laid  in  a  relatively  small  area.  In 
a  small  pool  near  the  main  Thornhill  pond  I  found,  on  Jan.  25, 1913,  59  masses 
in  an  area  approximately  10  X  15  feet.  Additional  masses  were  laid  in  this 
same  area  on  subsequent  nights.  Other  areas  in  this  same  pond  which  seemed 
to  be  of  exactly  the  same  character  were  not  utilized  by  the  species. 

"The  total  number  of  eggs  deposited  by  a  single  female  has  been  ascer- 
tained in  two  cases.  A  mated  pair  was  taken  from  the  Thornhill  pond  on 
January  31,  1914,  and  confined  in  an  aquarium.  Next  morning  a  total  of  730 
eggs  was  found.  The  average  number  of  eggs  in  a  clump  was  18.  Another  cap- 
tive which  began  laying  on  the  night  of  Mar.  7,  1922,  and  continued  up  until 
4  P.M.  of  the  day  following  deposited  a  total  of  1250  eggs;  the  average  of  the 
58  masses  laid  was  between  21  and  22.  The  largest  mass  contained  60  eggs  and 
there  were  two  with  the  minimum  of  9"  (T.  I.  Storer,  Calif.,  1925,  pp.  224- 

225)- 

"Insemination  occurs  at  the  moment  of  egg  extrusion.  The  male  brings  his 
cloacal  aperture  close  to  that  of  the  female,  discharges  a  quantity  of  trans- 
parent semen,  and  with  a  quick,  firm  extension  slides  his  feet  posteriorly  over 
the  sides  and  hips  of  the  female,  then  deftly  retracts  to  his  normal  position. 
Simultaneous  with  this  foot  action,  the  female  extrudes  a  clutch  of  eggs  into 


334  HANDBOOK  OF  FROGS  AND  TOADS 

the  cloud  of  sperm  about  her  cloaca.  Some  time  before  releasing  an  egg  mass 
the  female  often  scratches  at  the  surface  of  the  substratum  on  which  the  eggs 
are  to  be  deposited.  As  the  eggs  are  extruded  the  cloaca  is  brought  into  close 
contact  with  this  surface  and  attachment  is  thus  effected.  The  female  removes 
any  eggs  which  may  partially  adhere  to  the  cloaca  during  extrusion  by  a  pre- 
cise flexor-extension  reflex  of  the  hind  legs.  In  this  the  tarsi  are  applied  directly 
to  the  adhering  eggs  and  a  slow  extension  effects  the  removal. 

"Deposition  of  an  egg  mass  is  usually  followed  by  a  quiet  moment  during 
which  the  bodies  of  the  pair  become  slightly  more  inflated  than  normally. 
The  intervals  between  egg  deposition,  ranging  from  2  to  10  minutes  or  longer, 
are  spent  in  bursts  of  vigorous  activity,  mainly  on  the  part  of  the  female.  What 
function  this  may  serve,  if  any,  is  conjectural,  but  it  apparently  takes  place 
under  natural  as  well  as  artificial  conditions.  In  the  laboratory  such  periods 
are  followed  by  a  rest  and  then  further  oviposition.  Shortly  after  laying  is 
complete,  the  pair  becomes  separated.  The  entire  time  in  amplexus  has  been 
observed  to  range  from  8  to  40  hours  or  moie. 

''The  total  number  of  eggs  deposited  was  found  to  range  from  500  to  750; 
however,  Storer  reports  an  instance  of  1250  eggs  being  laid.  The  number  of 
eggs  per  clutch  is  usually  about  16,  but  varies  from  5  to  60"  (R.  E.  Smith,  Calif., 
1940,  pp.  379- ?8o). 

"Several  of  these  frogs  were  found  at  Big  Lake,  together  with  the  north- 
western toads.  Egg  clusters  in  the  litter  on  the  bottom,  near  the  edge  of  the 
lake,  were  in  jelly-like  masses  a  little  more  than  one  inch  in  diameter.  The 
egg  diameter  might  be  from  %a  to  Vf«  inch"  (F.  G.  Evenden,  Jr.,  Ore.,  1945, 
p.  252). 

"Hyla  regilla  also  spawns  abundantly  in  the  pool  on  Alpine  Road,  the  onset 
of  breeding  seeming  to  coincide  closely  with  that  of  A.  t.  caltforntense"  (V.  C. 
Twitty,  Calif.,  1941,  p.  2). 

"One  male,  Scaphiopus  /;.  intermontantts  (amplexus  inguinal)  when  col- 
lected was  clasping  a  female  Hyla  regilla  which  died  soon  after,  apparently 
from  rupture  of  the  abdominal  wall"  (G.  C.  Carl,  B.C.,  1942,  p.  129). 

Journal  notes:  Aug.  12,  1917.  We  camped  at  Jacumba,  Calif.,  beside  the 
Mexican  border.  ...  In  a  little  side  stream  of  the  creek  we  found  a  series 
of  H.  regilla  from  tadpole  to  transformation.  In  the  same  place  among  the 
weeds  of  the  moist  area  we  took  six  to  ten  adults  very  variable  in  color. 

Aug.  25.  In  Alta  Meadows,  we  found  no  end  of  transformed  Hyla  regilla. 
In  the  bog-ten  aced  pools  were  plenty  of  tadpoles  and  advanced  stages.  On 
the  trail,  R,  C.  Shannon  found  a  full-grown  frog  and  the  boys  reported  a 
large  one  from  Alta  Peak.  The  transformed  frogs  were  in  the  meadow 
land. 

Aug.  20-22,  1925,  Las  Vegas,  Nev.,  Tonapah  Road.  We  followed  one  tiny 
stream  back  into  the  field  and  returned  with  the  trophies  of  the  search:  6  Rana 
fisheri,  4  Hyla  regilla,  i  Bttjo  compactilis.  In  a  broad  springy  area  sedgy  and 


TREE  FROGS:  HYLIDAE  335 

shady,  we  picked  up  a  H.  regilla.  In  one  very  small  sedgy  area,  we  caught  four, 
three  of  which  had  triangles  between  the  eyes  and  some  dorsal  stripes.  A  half 
grown  one  was  green  with  no  triangle  and  only  the  vitta  back  of  the  eyes. 
Later  at  the  big  springs,  we  caught  one  that  looked  very  yellow  with  very  in- 
distinct pattern.  We  caught  several  tadpoles. 

Starting  in  January,  1942,  we  traveled  northward  west  of  the  mountains 
from  southern  California  to  Washington  and  southward  east  of  the  moun- 
tains, arriving  in  Nevada  in  mid-May.  The  Journal  notes  of  that  period  give 
a  specimen  cross  section  of  the  activities  of  this  widespread  species. 

Jan.  31,  1942.  Stayed  at  Julian,  Calif.,  for  night.  Last  night  heard  a  chorus 
of  Hyla  regilla.  The  early  season  and  the  chorus  each  make  me  think  of 
Pseudacris  choruses  in  early  spring.  It  is  so  unlike  most  true  Hyla  barks  of 
the  U.S.A. 

Feb.  i.  Went  to  Cuyamaca.  In  Julian  region  where  I  heard  the  choruses  last 
night  took  about  seven  or  eight  male  H.  regilla.  One  female  and  one  very 
small  one.  Tonight  the  male  is  mated  axillary  fashion.  Tonight  heard  a  large 
chorus  of  H.  regilla  but  no  others. 

Feb.  7.  Five  miles  out  of  Monterey  at  12:30  P.M.,  7o°F.,  sunlight.  We  heard 
chorus  of  H.  regilla  in  an  overflow  grassy  area  near  base  of  pine  hill. 

Feb.  8.  Salinas  to  Castroville,  Calif.  Could  find  no  Hyla  regilla  eggs.  Do 
they  lay  them  in  dense  vegetation  at  the  edge  of  water  like  P.  feriarum  of  Car- 
lisle, Pa.? 

Feb.  12.  Visited  W.  M.  Ingram.  He  had  two  Hyla  regilla  masses  of  eggs 
ready  to  hatch. 

Feb.  14.  Trip  to  R.mcho  del  Oso — Prof.  Theodore  S.  Hoover  owner.  Found 
H.  regilla  in  swampy  area.  Up  the  creek  to  sawmill.  Here  in  a  little  pool  found 
very  young  H.  regilla  tadpoles.  Rodgers  took  a  large  H.  regilla  with  reddish 
down  middle  of  back. 

March  8.  Stanford  artificial  lake.  Found  in  an  overflow  shallow  some  eight 
fresh  masses  oi  H.  regilla  eggs.  The  yellowish  underside  is  evident.  Mass  an 
inch  or  less  in  diameter.  Sometimes  two  masses  on  one  grass  blade.  One  or 
two  masses  hatching.  Found  some  tadpoles  VL>  inch  long. 

March  17.  Later  went  north  of  Crescent  City  on  Route  101  about  5  miles.  In 
a  pond,  countless  tads  of  H.  regilla  and  a  few  egg  masses  almost  hatching. 

March  18.  Took  Hyla  regilla  tads  Carpcntcrville,  Ore.  H.  regilla  calling  at 
midday. 

March  20.  Up  McKenzie  River  watershed  with  Miss  Ruth  E.  Hopson  of 
Springfield,  Ore.  In  a  pond  with  fallen  logs,  no  end  of  H.  regilla  migrating 
to  the  pond,  mostly  females.  1800-1900  elevation. 

March  26.  Started  from  Aberdeen  at  9  A.M.  Foggy.  At  Springfield  School 
Dist.  No.  12  saw,  beside  road,  ponds  with  pussy  willows  in  them.  Here  were 
Hyla  regilla  masses,  some  just  hatching.  At  Humptulips  River  crossing,  in 
pools,  caught  a  Hyla  regilla.  Two  miles  farther  took  a  female. 


336  HANDBOOK  OF  FROGS  AND  TOADS 

March  27.  Hoh  to  Borachiel  rivers.  In  meadow  heard  one  lone  H.  regilla. 
Saw  here  an  egg  mass  of  H.  regilla. 

March  28.  Went  to  Olympic  Park  headquarters.  Mr.  Macy  gave  me  a  permit 
and  a  map.  In  their  pond  saw  several  fresh  egg  masses  and  some  about  to 
hatch  (H.  regilla).  Just  as  we  turned  on  to  Elwah  road,  in  one  meadow  pond 
saw  some  500-800  egg  masses  of  H.  regilla  hatching  and  fresh.  Anna  caught 
a  pair. 

April  4.  Just  out  of  Maupin,  Ore.,  to  the  south  of  town  in  a  gravel-pit  pool 
saw  some  six  or  eight  H.  regilla  masses.  One  H.  regilla  jumped  in. 

April  5.  Lapine,  Ore.,  roadside  ditch.  The  H.  regilla  are  calling  loudly  but 
could  find  no  egg  masses.  Flatland,  open,  some  cultivated,  some  meadows, 
trees. 

April  10.  From  Medford  to  Project  City,  in  rather  barren  stream  took  some 
almost  full-grown  H.  regilla  tadpoles. 

April  ii.  Chico,  Calif.,  in  a  spring  watering  trough  were  large  H.  regilla 
tadpoles. 

April  12.  Out  with  Tracy  and  Ruth  Storer.  Between  Michigan  Bar  and 
Consumne  spotted  beside  road  a  possible  A.  t.  cahforniense  pond.  H.  regilla 
tads  mature.  White  egret  here  feeding. 

April  21.  Below  Tollhouse,  Calif.,  in  one  pond  we  took  two  H.  regilla  but 
no  R.  b.  sierrae. 

April  23.  Turned  toward  the  coast  again  to  Atascadero,  Calif.  Heard  no 
end  of  H.  regilla  last  night. 

May  i.  In  French  Valley,  Calif.,  in  drying  pool  took  H.  regilla  tads,  one 
adult.  At  a  culvert  took  H.  regilla  tads  mature  and  one  transforming.  No  end 
of  Brewers  black  birds  around  this  pool  of  tadpoles  and  aquatic  insects. 

May  5.  Three  and  a  half  to  four  miles  south  of  Buckman  Springs,  Calif.,  at 
bridge  below  confluence  La  Posta  and  Cottonwood  Cr.,  took  two  H.  regilla. 
Saw  no  H.  regilla  tads. 

May  14,  Springdale,  Nev.  Bobbie  Shoshone  went  out  last  night  and  caught 
one  Bufo  b.  nelsom  and  several  Hyla  regilla  (3). 

May  15,  Las  Vegas,  Nev,  Went  out  Tonopah  road  25  miles  and  returned 
after  dark.  Near  trailer  camp  was  a  large  pond  with  bullfrogs  in  it  and  near 
by  a  big  chorus  of  Hyla  regilla. 

Authorities'  corner: 

F.  C.  Test,  Gen.,  1899,  2I>  $*•  J-  Grinnell,  J.  Dixon,  J.  M.  Linsdale, 

W.  P.  Taylor,  Nev.,  1912,  p.  343.  Calif.,  1930,  p.  141. 

J.  Van  Denburgh  and  J.  R.  Slevin,  L.C.,     A.  and  R.  D.  Svihla,  Wash.,  1933,  p. 

1914,  p.  135.  127. 

A.  G.  Ruthven  and  Helen  T.  Gaige,     S.  G.  Jewett,  Jr.,  Ore.,  1936,  p.  71, 

Nev.,  1915,  p.  15. 


TREE  FROGS:  HYLIDAE 


337 


Plate  LXX.  Hyla  regilla.  1,2,6.  Females  Plate  LXXI.  Hyla  septentrionalis  (X%). 

(XO-  3»4-  Mai*  (X%)-  5-  Female  (X%).         1-3-  Females. 


338  HANDBOOK  OF  FROGS  AND  TOADS 

Giant  Tree  Frog 

Hyla  septentnonalis  Boulenger.  Plate  LXXI;  Map  19. 

Range:  On  Oct.  30,  1931  (Fla.,  p.  140),  Dr.  T.  Barbour  placed  it  as  estab- 
lished at  Key  West,  Fla.  In  1914  (p.  347)  he  had  it  from  New  Providence, 
Andros  Island,  Rum  Cay,  Cuba,  and  Grand  Cayman.  Matecumbe,  Fla.,  and 
farther  northward. 

Habitat:  Cisterns,  wells,  outbuildings,  drain  pipes;  banana  trees,  large  leaf 
axils. 

"Old  cisterns  and  damp  out-buildings;  in  the  axils  of  palm  and  Caladium 
leaves;  banana  trees;  Barbour  (1931)  records  it  from  drain  pipes.  Abundance: 
Locally  common;  I  have  seen  twenty-five  or  thirty  clinging  to  the  walls  of  a 
single  cistern  in  Key  West.  Habits:  Chiefly  nocturnal,  but  often  calling  and 
foraging  during  the  day  in  the  dilapidated  cisterns  which  abounded  on  the 
island  before  'New  Deal'  improvements.  During  humid  weather  they  may  be 
found  in  trees  and  shrubs  some  distance  from  water.  I  have  discussed  septen- 
trionalis  with  some  of  Key  West's  most  venerable  citizens  and  have  found 
none  who  can  recall  a  time  when  the  blatant  choruses  of  the  'bull  frogs'  were 
not  to  be  heard  after  summer  rains.  Feeding:  They  often  congregate  around 
street  lights  at  night;  I  have  watched  them  feeding  on  cockroaches  and  on 
lepidopterous  larvae;  they  are  batracophagous  and  cannibalistic  in  captivity" 
(A.  F.  Carr,  Jr.,  Fla.,  1940^  p.  62). 

Size:  Adults,  2%-^%  inches  (64-130  mm.),  "There  is  a  great  size  disparity 
in  the  breeding  males;  individuals  that  I  have  measured  varied  between  40 
and  73  mm."  (A.  F.  Carr,  Jr.,  Fla.,  1940,  p.  62). 

In  our  Matecumbe  Key  series  (March  18, 1934)  we  took  males  48,  48,  49,  49, 
50,  50,  51,  52,  52.5,  57,  58,  59, 60,  and  61  mm.  in  length,  fourteen  in  all,  and  one 
small  female,  52.5  mm.  In  1940  at  Key  West,  Mr.  and  Mrs.  R.  H.  McCauley, 
May  5,  secured  two  males,  46  and  48  mm.,  and  one  large  female,  82  mm.;  on 
April  10,  1940,  they  secured  on  Stock  Island,  Monroe  Co.,  Fla.,  two  females 
75.5  and  77  mm.  in  length.  The  largest  we  have  is  from  Key  West,  taken  by 
M.  K.  Brady  and  O.  C.  Van  Hyning.  It  is  a  female,  92  mm.  in  length.  We 
have  seen  large  specimens  but  recall  none  over  4  inches.  With  our  limited 
series  we  can  only  approximate  the  sex  sizes  as  follows:  males,  46-75  mm.; 
females,  52.5-130.0  mm. 

General  appearance:  This  is  a  large  tree  toad.  The  head  is  broad;  the  out- 
line of  the  skull  evident,  as  the  skin  is  united  to  the  skull;  canthus  rostralis 
and  nostrils  very  prominent.  The  top  of  the  head  is  smooth;  the  eyelids  and 
back  are  roughened  with  large  and  fine  tubercles.  The  most  conspicuous  char- 
acters are  the  very  large  disks  on  fingers  and  toes,  those  on  the  fingers  being 
fully  as  large  as  the  tympanum.  The  eyes  are  large  and  prominent,  the  iris 
with  brilliant  orange  tints.  The  color,  when  the  frog  has  been  under  cover, 
becomes  a  dull  olive-green,  but  in  the  light  becomes  citrine,  turtle  green,  or  oil 


TREE  FROGS:  HYLIDAE 


339 


yellow,  with  indistinct  dorsal  spots  of  dull  citrine  or  grayish  olive.  The  legs 
are  barred  with  the  same.  The  rear  of  the  ftmur  is  reticulated  with  the  same. 
The  throat  is  pale,  huffy,  and  slightly  granular;  the  rest  of  the  venter  is  con- 
spicuously and  roughly  granular  and  dull  yellow  in  color,  with  the  underside 
of  the  femur  a  deeper  yellow,  and  the  axilla  bright  yellow  with  a  wash  of  the 
same  color  along  the  sides.  The  tubercles  under  the  joints  of  feet  and  hands 
are  prominent  and  pointed. 

"Male  with  two  external  vocal  vesicles,  each  being  situated  near  the  angle  of 
the  mouth;  during  the  breeding-season  the  inner  side  of  the  first  finger  cov- 
ered with  blackish  rugosities.  From  snout  to  vent  75  millim."  (G.  A.  Bou- 
lenger,  Gen.,  1882,  pp.  368-369). 

Color:  Female.  Key  West,  Fla.,  from  M.  K.  Brady  and  O.  C.  Van  Hyning, 
May  5, 1932.  Trough  of  casque  and  concave  lorcal  region  sulphur  yellow,  ecru- 
olive,  or  buffy  citrine.  Postorbital  horns  of  casque  pale  turtle  green  or  oil 
yellow.  Wash  of  this  color  i  inch  or  more  wide  down  middle  of  back.  Area 
below  and  behind  eye  to  below  tympanum,  upper  parts  of  fore  limbs,  and 
throat  light  buff  or  pale  ochraceous-buff.  The  sides  of  body  with  slight  wash 
of  the  same  color.  From  axilla  along  either  side  of  belly  to  groin  is  a  wash 
of  wax  yellow  or  primuline  yellow  in  axil.  Two  bars  on  thighs  ceasing  halfway 
as  they  meet  reticulations  on  rear  of  thigh.  Tibia  with  two  complete  bars.  Bars 
and  obscure  spots  on  the  rear  half  of  back  are  dull  citrine,  grayish  olive,  or 
buffy  citrine.  The  dark  reticulations  of  the  thighs  are  colored  like  bars  of 
thighs  with  the  interspaces  green  yellow.  There  is  a  little  of  same  color  on 
front  of  femur  and  in  groin.  The  top  of  the  foot  has  decided  wash  of  same 
color  as  top  of  fore  limbs.  The  digits  are  turtle  green.  The  center  of  tympanum 
like  back  surrounded  by  ring  of  mars  brown.  The  iris  is  capucine  yellow  or 
ochraceous-buff  finely  dotted  with  black.  Sometimes  the  iris  has  considerable 
of  the  color  of  the  back  or  of  dorsum  of  fore  limbs.  Undersides  of  fingers  and 
toes  are  cinnamon-drab, 

Structure:  Tongue  subcircular;  head  broader  than  long  in  adults;  casque 
emarginate  behind;  snout  rounded,  contained  two  times  in  head  to  tympa- 
num; pollex  rudiment  not  free  projecting;  disks  conspicuous;  no  interocular 
bar;  upper  eyelid  small;  tympanum  distinct,  %-%  diameter  of  eye.  Dunn  in 
1937  (Copeia,  p.  166)  placed  H.  septentrtonalis  in  the  Hyla  bmnnca  group. 

"Fingers  slightly  webbed;  no  projecting  rudiment  of  pollex;  toes  two-thirds 
webbed;  disks  of  fingers  nearly  as  large  as  the  tympanum,  of  toes  smaller; 
subarticular  tubercles  well  developed;  a  slight  fold  along  the  inner  edge  of 
the  tarsus.  The  hind  limb  being  carried  forwards  along  the  body,  the  tibio- 
tarsal  articulation  reaches  between  the  eye  and  the  tip  of  the  snout  .  .  ." 
(G.  A.  Boulenger,  Gen.,  1882,  pp.  368-369). 

Voice:  "This  sound  is  like  the  jerky  pulling  of  a  rope  through  an  unoiled 
pulley  and  is  very  characteristic"  (T.  Barbour,  Fla.,  1931,  p.  140). 

"The  choruses  arc  unique  in  their  heterogeneity  of  pitch  and  timbre,  The 


340  HANDBOOK  OF  FROGS  AND  TOADS 

call  is  a  rasping  snarl  somewhat  comparable  to  that  of  Scaphiopus 
The  pitch  of  non-musical  sound  is  difficult  to  determine,  but  I  would  judge 
that  the  individual  notes  vary  through  at  least  one  octave"  (A.  F.  Carr,  Jr., 
Fla., 1940^  pp.  62-63). 

We  were  first  attracted  to  them  at  Upper  Matecumbe  Key  by  a  huge  chorus 
near  our  cabin.  In  the  circle  of  bushes  around  a  small  very  deep  pond  were 
these  gaint  tree  frogs.  We  neglected  to  characterize  their  call,  to  photograph 
them,  or  to  describe  them.  Doubtless  there  were  eggs,  but  Elaphe  rosacea  was 
our  chief  interest  at  the  moment. 

Breeding:  "June  I4  to  September  16  (this  period  corresponds  suspiciously 
with  the  academic  summer-vacation  season;  however  I  looked  for  them  in 
vain  during  the  week  of  December  18-23,  1932).  The  eggs  are  laid  in  tempo- 
rary drainage  ditches,  in  the  flooded  basements  of  wrecked  buildings,  and  in 
cisterns.  The  young  emerge  at  15.5-16.5  mm.  Before  the  tail  is  absorbed,  the 
back  becomes  granular  and  warty  as  in  the  adult;  the  immature  differs  strik- 
ingly from  the  adult  in  the  possession  of  a  dorso-lateral  white  stripe  extending 
from  the  hind  margin  of  the  orbit  to  the  groin.  E.  Lowe  Pierce  and  I  found 
all  stages,  from  newly  hatched  larvae  to  breeding  adults,  in  an  old  cistern, 
June  16, 1934.  We  found  a  tremendous  number  (300-400)  of  recently  emerged 
individuals  perched  on  the  twigs  of  a  small  thorny  bush  in  a  vacant  lot,  the 
night  of  June  14, 1934"  (A.  F.  Carr,  Jr.,  Fla.,  1940^  p.  62). 

March  18,  1934,  Matecumbe  Key,  Fla.  Males  have  enlarged  thumbs;  on 
some  smaller  males  the  horny,  sharp-edged  ridge  is  not  so  black.  In  general,  it 
is  a  dark,  horny  excrescence  on  inner  side  and  upper  side  of  the  the  thumb;  it 
is  semilunar,  extending  from  base  of  thumb  to  base  of  last  segment  of  thumb. 
Outer  edge  acute  and  making  a  sharp  acute  edge,  apparent  from  ventral  side 
of  thumb.  It's  an  expanded  addition  or  prepollex  on  side  of  the  thumb 
(pollex). 

Some  of  these  males  are  almost  uniform,  some  streaked  in  groin  like 
H.  batidmii,  some  very  mottled  on  back;  almost  all  with  leg  bands,  some  nar- 
row, some  wider.  Several  with  two  prominent  stripes  extending  backward 
from  eyes. 

May  5,  1940.  Dr.  and  Mrs.  R.  H.  McCauley  took  one  specimen  just  beyond 
transformation,  40  mm.  in  length  (with  24-mm.  body  and  remnant  of  tail 
16  mm.).  This  specimen  has  the  tail  crests  and  musculature  heavily  mottled 
with  black.  They  also  took  the  following  interesting  series:  at  transformation 
— 21  mm.;  past  transformation — 25, 26,  28, 30,  30,  32,  33,  36,  37  mm.;  male  with 
black  on  thumb — 46,  48  mm. 

Key  West,  Fla.  "The  largest  of  fourteen  specimens  collected  was  a  female 
measuring  92  mm.  in  length  of  body,  and  210  mm.  stretched  out.  Eggs,  tad- 
poles, and  recently  transformed  frogs  were  also  collected"  (E.  R.  Allen  and 
R.  Slatten,  Fla.,  1945,  p.  25). 

Journal  notes:  March  18,  1934,  Upper  Matecumbe  Key,  Fla.  Came  to  Cari- 


TREE  FROGS:  HYLIDAE  34! 

bee  Colony.  At  night  heard  a  great  chorus  near  the  water  tank.  There  were 
countless  small  and  half-grown  Hyla  septentnonalis  in  the  pond.  No  end  of 
Hyla  sqttirdla  calling.  Are  H.  squirella  the  prey  of  the  giant  tree  frog?  From 
limbs  higher  than  my  head  big  tree  frogs  would  leap  into  the  middle  of  the 
pond  with  a  pronounced  ^erplunf^  (like  a  toad).  In  looking  for  the  frogs 
Anna  espied  in  the  bushes  a  fine  Elaphe  rosacea.  Does  it  feed  on  the  frogs  ? 

Authorities'  corner: 
G.  A.  Boulenger,  Gen.,  1882,  pp.  368-     T.  Barbour,  Fla.,  1914,  p.  238. 

369.  T.  Barbour,  Fla.,  1931,  p.  140. 

L.  Stejneger,  Fla.,  1905,  p.  330.  C.  S.  Brimley,  N.C.,  1940,  p.  23. 

Squirrel  Tree  Frog,  Southern  Tree  Frog,  Southern  Tree  Toad,  Scraper  Frog, 
Rain  Frog,  Squirrel  Tree  Toad,  Squirrel  Hyla 

Hyla  squirella  Latreille.  Plate  LXXII;  Map  24. 

Range:  Virginia  to  Texas,  and  north  up  the  Mississippi  basin  to  Indiana. 
Near  Columbia,  S.C.,  Corrmgton  (S.C.,  1929,  p.  66)  "found  them  on  the  fall 
line  but  never  in  good  typical  piedmont  territory." 

Habitat:  In  and  around  buildings;  about  wells;  in  bushes,  trees,  or  vines; 
in  fields  and  gardens.  It  breeds  in  open  ponds  in  the  pine  barrens,  or  in  shallow 
roadside  pools. 

"This  species  was  found  on  the  ground  about  shady  hammock-land  on  Boca 
Chica  Key.  I  also  saw  one  on  a  Gumbo  Limbo  tree  on  Vaca  Key"  (H.  W. 
Fowler,  Fla.,  1906,  p.  109). 

"Common  when  breeding;  frequents  gardens  and  is  often  seen  in  crannies 
about  porches  of  houses"  (O.  C.  Van  Hyning,  Fla.,  1933,  p.  4). 

"Shows  little  discrimination  in  major  habitat  selection.  It  exhibits  some 
preference  for  the  more  open  wooded  areas  such  as  high  pine  and  mixed  ham- 
mock and  for  edificarian  situations.  Abundance. — The  commonest  Florida 
Hyla"  (A.  F.  Carr,  Jr.,  Fla.,  1940^  p.  61). 

Size:  Adults,  %-i%  inches.  Males,  23- $6  mm.  Females,  23-37  mm. 

General  appearance:  This  species  is  small,  delicate,  and  with  smooth  skin. 
The  head  is  short,  eyes  prominent  with  black  pupil  and  bronzy  iris.  The  back 
is  green  or  brownish  in  color  with  at  least  a  partial  transverse  bar  between  the 
eyes  and  with  white  on  the  upper  lip.  Frequently  there  are  rounded  spots  on 
the  back.  The  rear  of  femur  is  not  spotted.  There  is  a  light  line  below  the  eye 
and  over  the  shoulder.  There  may  be  a  light,  irregular  line  along  the  side  just 
above  the  belly. 

"The  colours  of  this  animal  are  even  more  changeable  than  in  any  species 
with  which  I  am  acquainted.  I  have  seen  it  pass  in  a  few  moments  from  a  light 
green,  unspotted  and  as  intense  almost  as  that  of  Hyla  lateralis,  to  ash  colour, 
and  to  a  dull  brown  with  darker  spots;  the  spots  also  at  times  taking  on  dif- 
ferent tints  from  the  general  surface.  The  markings,  too,  vary  exceedingly  in 


34* 


HANDBOOK  OP  FROGS  AND  TOADS 


different  individuals,  the  white  line  on  the  upper  lip  and  the  band  between 
the  orbits  alone  are  constant"  (J.  E.  Holbrook,  Gen.,  1842,  p.  124). 

See  M.  C.  Dickerson,  Gen.,  1906,  pp.  149-150  for  a  discussion  of  the  wonder- 
ful change  of  color  in  this  form.  For  variability  of  color  in  Hyla  squirella  and 
Hyla  ctnerea,  see  our  "Journd  notes"  from  Pass  Christian  and  Bay  St.  Louis 
under  Hyla  cinerea.  We  have  not  seen  Bryan  P,  Glass's  (Tex.,  1946,  pp.  101- 
103)  specimens  of  "A  New  Hyla  from  South  Texas"  but  suspect  he  has  Hyla 


Map  24 

squirella  in  one  of  its  variable  disguises  in  his  new  Hyla  flavigula.  Of  course 
examination  of  specimens  alone  can  confirm  this  impression.  Hyla  squirella 
adult  males  and  females  range  as  low  as  23  mm.  and  as  high  as  37  mm.  Glass's 
specimens  may  be  about  the  mean  of  the  squirrel  tree  frog.  This  species  has 
been  recorded  four  times  inland  from  Aransas  country  and  north  of  the 
Nueces  River,  Tex.  P.  H.  Pope  (Tex.,  1919)  found  H.  squirella  the  most 
abundant  Hyla  of  the  Houston  area. 

Color:  Males.  June  22, 1922.  Back  citrine  or  buffy  citrine  in  one;  on  another 
ecru-olive  or  light  yellowish  olive;  rear  back  and  Lop  of  hind  legs  chrysolite 
green  to  lime  green.  Under  parts  of  hand,  also  hind  foot,  orange-rufous,  xan- 
thine  orange,  to  orange  chrome;  hind  limb  posterior  and  anterior  faces  and 
throat  raw  sienna  or  mars  yellow;  throat  of  another  aniline  yellow;  in  another 
hind  leg  parts  raw  sienna.  Stripe  on  upper  jaw  to  body  greenish  yellow  or 


TREE  FROGS:  HYLIDAE  343 

lemon  yellow.  Belly  cream  color  or  ivory  yellow.  Tympanum  hazel  or  russet. 
Iris  spotted  black  and  army  brown — fawn  in  one,  deep  chrome  and  antique 
brown  in  another.  Greenish  on  either  side  of  throat  but  not  clearly  defined  or 
definite  as  in  Hyla  andersonii  and  Hyla  cinerea  males.  One  male  held  in  the 
hand  in  good  light  became  courge  green  and  another  oil  green.  A  small  last 
year's  H.  squirella  was  light  dull  green-yellow  above,  and  upper  jaw  stripe 
light  chalcedony  yellow  or  pale  chalcedony  yellow.  Sometimes  those  in  water 
may  be  greenish,  while  those  on  edges  or  on  land  may  be  brownish. 

Females.  July  3,  1922.  Four  females  of  four  different  pairs  had  under  parts 
white,  with  no  discolored  throats.  Throat  slightly  primrose  yellow,  reed  yel- 
low, or  white.  Fore  limbs,  groin,  hind  limbs  before  and  aft,  tibia,  and  hind 
feet  reed  yellow  to  olive-yellow.  In  one,  yellow  ocher  fore  and  rear  part  of 
femur,  underside  of  tibia,  and  foot  and  fore  foot.  The  females,  unless  very 
large,  have  none  of  the  bright  orange-rufous,  xanthine  orange,  or  orange 
chrome  of  the  males. 

Structure:  Form  delicate;  skin  smooth;  canthus  rostralis  well  marked,  but 
not  sharply  edged;  throat  of  male  raw  sienna  or  yellow  with  light  area  of 
greenish  on  either  side;  vocal  sac  a  large  hyaline  subgular  pouch. 

Voice:  This  call  is  a  harsh  trill,  regular,  mostly  continuous,  15  calls  in  10 
seconds,  but  not  very  loud.  It  could  not  be  heard  a  few  rods  away.  One  gave 
67  pumps  in  45  seconds.  "Call  rasping,  but  not  very  coarsely  so;  rather  ventrilo- 
quistic;  flat;  short  to  Italian  a;  aaaa  or  waaaaal^'  (A.  F.  Carr,  Fla.,  1934,  p.  22). 

Another  great  chorus  came  Aug.  16  and  17  after  a  half-day's  ram  when 
about  2  inches  of  rain  fell,  maxima  82-91°,  minima  65-74°.  Spadefoots  and 
Rana  capito,  subterranean  species,  came  out  to  breed  and  Hyla  squirella  as 
well. 

In  general  this  species  calls  even  by  day  in  rain  or  before  an  imminent  rain 
(July  9) .  After  a  downpour  of  a  warm  rain  1-5  inches  they  became  very  active. 
As  every  observer  has  remarked,  heavy  warm  rains  bring  them  out  in  mid- 
summer. Our  latest  record  of  calling  came  September  17. 

R.  F.  Deckert  (Fla.,  1915,  p.  3)  at  Jacksonville,  Fla.,  described  its  cry  as  fol- 
lows: "The  cry  is  rather  coarse,  sounding  like  'era,  era,  era/  etc.  with  a  second's 
interval  between  each  note."  The  same  author  (Fla.,  1921,  p.  22)  in  Dade 
County,  Florida,  wrote,  "Their  rasping  calls  were  heard  in  May  and  June 
at  igth  Street,  Miami,  at  Donn's  Nursery  and  at  22nd  Street  in  company.  .  .  ." 

Breeding:  This  species  breeds  from  April  to  August.  The  eggs,  single  on 
the  bottom  of  shallow  pools,  are  brown  above  and  cream  below,  with  outline 
distinct  and  jelly  firm.  The  egg  is  %o-%s>  inch  (0.8-1.0  mm.),  the  inner  en- 
velope ^o-Vis  inch  (1.2-1.6  mm.),  the  outer  %o-M2  inch  (1.4-2.0  mm.). 
The  egg  complement  is  950.  The  citrine  drab  tadpole  is  small,  i/4  inches 
(32  mm.),  its  tail  long,  the  tail  tip  acuminate  with  a  flagellum.  The  tooth 
ridges  are  %.  After  40  to  50  days,  the  tadpoles  transform  from  June  to  Sep- 
tember at  %G  inch  (11-13  mm.). 


344  HANDBOOK  OF  FROGS  AND  TOADS 

"I  have  records  from  April  2  to  August  20.  Full  choruses  come  with  July 
electric  storms.  The  males  usually  sing  in  water  from  half  an  inch  to  an  inch 
deep  at  the  edges  of  temporary  pools  or  ditches  or  of  temporarily  inundated 
portions  of  pond  margins.  The  choruses  are  large.  Wright  (1933)  says  that 
the  call  is  not  very  loud;  it  has  little  carrying  power,  but  at  close  range  a  large 
chorus  assaults  the  ear-drums  mercilessly"  (A.  F.  Carr,  Jr.,  Fla.,  1940!),  p.  61). 

Journal  notes:  We  found  these  frogs  on  porches,  in  chinaberry  trees,  oaks, 
and  other  trees,  as  well  as  in  vines  around  the  houses,  in  fields  and  gardens 
around  buildings,  in  open  ponds,  in  pine  barrens,  in  pine  and  oak  groves, 
along  roads,  and  in  shallow  roadside  and  pine  barren  pools.  On  June  22, 1922, 
a  boy  at  Camp  Pinckney,  Ga.,  brought  me  a  Scraper  taken  in  his  own  house. 

July  3, 1922.  Along  the  Folkston  road,  Ga.,  in  temporary  pools  and  ditches 
with  Btifo  terrestris  were  plenty  of  Hyla  squirella.  There  is  more  vibration  in 
the  call  of  Hyla  jemoralis.  I  could  not  hear  the  Hyla  squirella  a  few  rods 
away.  The  calls  of  Hyla  femoralis  and  Bttfo  quercicus  drown  it  out.  Hyla 
squirella  does  sometimes  croak  from  the  water  surface  when  sprawled  on  the 
water. 

Okefinokee  Swamp,  Ga.  On  July  3,  1922,  about  11:55  P.M.,  we  were  at 
Anna's  pond.  We  heard  the  Hyla  squirella  at  a  distance  between  two  distant 
houses.  In  the  saw  palmettos  and  the  grass  stools  were  many  scrapers.  We 
found  one  pair  in  the  grass  near  the  edge.  Others  were  found  in  saw  palmettos 
about  the  border  of  the  pool.  The  same  night,  near  Trader's  Hill,  we  came 
to  a  grassy  overflow  pond.  In  a  clump  of  bushes  and  saw  palmetto  were  sev- 
eral Hyla  squirella.  In  grassy  stools  in  shallow  water  were  others. 

On  August  n,  1922,  at  Camp  Pinckney,  Ga.,  2-3  P.M.,  we  heard  several. 
At  8:30  we  returned  to  Camp  Pinckney.  There  were  countless  Hyla  squirella 
males  on  the  ground  in  a  road  filled  with  temporary  pools,  in  water  1-3  inches 
deep.  The  vocal  sac  is  hyaline,  more  or  less  inflated  for  some  time.  The  call 
is  not  so  fast  as  that  of  Hyla  jemoralis,  but  swift  nevertheless.  Those  in  water 
were  greenish,  those  at  the  edges  of  the  pool,  brownish.  .  .  .  We  could  find 
no  females.  Some  males,  though  quite  small,  were  croaking. 

Authorities1  corner: 

W.  Bartram,  Gen.,  1791,  p.  278.  A.  F.  Carr,  Jr.,  Fla.,  1940^  p.  61. 

P.  H.  Pope,  Tex.,  1919,  pp.  96-97.  G.  L.  Orton,  La.,  1947,  pp.  369-375. 

A.  H.  Wright,  Ga.,  1932,  p.  312. 

Common  Tree  Toad,  Common  Tree  Frog,  Tree  Toad,  Northern  Tree  Toad, 
Changeable  Tree  Toad,  Chameleon  Hyla,  Varying  Tree  Toad 

Hyla  versicolor  versicolor  (Le  Qonte).  Plate  LXXIII;  Map  20. 

Range:  Maine,  southern  Canada,  west  to  Manitoba,  Minnesota,  South  Da- 
kota, south  through  Kansas,  Oklahoma,  to  the  Gulf  States  and  northern  Flor- 
ida (Texas  and  Arkansas  in  part  only). 


TREE  FROGS:  HYLIDAE 


345 


LXX/7.  #y/0  squirella  (XO- 
Male  croaking.  2,3,5.  Males.  4.  Female. 


LXX//7.  Hy/0  versicolor  versicolor. 
1,2,6.  Males  (X%)-  3-  Female  (X%)-  4- 
Males  croaking  (X%).  5-  Eggs  (X%). 


346  HANDBOOK  OF  FROGS  AND  TOADS 

Habitat:  Trees,  mossy  or  lichen-covered  stone  fences,  decaying  fruit  trees. 

Size:  Adults,  iJ/4-2%  inches.  Males,  32-51  mm.  Females,  33-60  mm. 

General  appearance:  This  frog  varies  in  color  from  pale  brown  to  ashy  gray, 
to  green;  it  has  a  granular  skin,  dark  irregular  star  on  upper  part  of  the  back, 
a  black  bar  on  the  upper  eyelid,  black-bordered  green  bars  on  the  legs,  a  black- 
bordered  light  spot  below  each  eye.  It  might  easily  be  taken  for  a  stone  or  bit 
of  bark  with  lichen  on  it.  The  rear  of  the  femur  has  dark  reticulations  on 
orange.  The  groin,  axilla,  and  under  parts  of  hind  limbs  are  orange. 

Color:  Male.  Belly  and  lower  breast  white  or  cartridge  buff.  Breast  between 
fore  limbs,  which  is  the  vocal  sac,  pale  flesh  color.  Middle  of  throat  dull  citrine 
or  ecru-olive  with  black  spots.  Chin  like  belly  with  blackish  spots.  Underside 
fore  limbs  like  belly.  Underside  of  hind  legs  and  rear  of  thigh  bright  orange 
or  orange-rufous.  The  spots  on  the  back,  one  of  which  is  an  irregular  star  on 
upper  back,  are  citrine-drab  or  deep  olive  with  black  borders. 

Female.  Raleigh,  N.C.,  from  A.  P.  Chippey,  May  12,  1950.  Under  parts  in- 
cluding throat  white  becoming  almost  hyaline  on  underside  of  femur.  Front 
of  femur,  groin,  inner  side  of  tibia,  tarsus  and  first  toe,  and  axilla  cadmium 
yellow,  deep  chrome,  or  in  places  mars  yellow.  Rear  of  femur  raw  sienna,  with 
numerous  spots  of  cadmium  yellow  or  light  cadmium.  Back  of  forelegs  and 
hind  legs  and  sides  pale  olive  gray,  pearl  gray,  or  light  mineral  gray,  becoming 
on  back  tea  green  or  grayish  olive  or  deep  olive-buff.  Star  or  spot  in  middle  of 
upper  back  is  vetiver  green  or  grayish  olive  with  dark  grayish  olive  border. 
Two  bars  on  femur,  tibia,  and  forearm  olive  gray  to  mineral  gray  with  dark 
borders.  Oblique  bar  across  eyelid  of  each  eye  but  not  meeting  on  meson  is 
same  color  as  star  spot  of  back.  Tympanum  drab  or  cinnamon-drab.  Spot  be- 
low eye  not  so  prominent  as  in  male  described,  is  olive-buff.  Iris  black  with 
ecru-drab  or  light  vinaceous-fawn  spots.  Iris  rim  avellaneous  or  vinaceous 
fawn;  iris  rim  broken  in  front,  below,  and  somewhat  behind. 

Structure:  Skin  rough,  warty;  conspicuous  disks  on  fingers  and  toes. 

Voice:  The  call  is  a  loud,  resonant  trill,  ending  abruptly,  10  or  n  calls  in 
half  a  minute.  About  the  middle  of  May,  at  Ithaca,  they  are  in  the  chorus  stage. 
In  the  evening,  all  over  the  University  hill  and  the  hills  near  by,  along  the 
wooded  ravines,  in  the  thickety  edges  and  woods  of  our  marshes,  we  may 
stumble  upon  the  noisy  tiee  toads  slowly  approaching  the  nearest  breeding 
place.  In  one  instance  their  resort  is  a  pond  at  the  end  of  a  long  hedge.  Here, 
at  the  breeding  season,  every  evening  and  sometimes  after  a  thundershower 
by  day,  the  males  can  be  heard  all  along  its  length,  slowly  bound  for  the  one 
objective,  the  pool,  where  some  have  already  arrived. 

Breeding:  These  tree  toads  breed  from  the  end  of  April  to  Aug.  n.  The 
brown  and  cream  or  yellow  eggs  are  laid  in  small  scattered  masses  or  packet; 
of  not  more  than  30  to  40  eggs  on  the  surface  of  quiet  pools,  the  packets  loosel] 
attached  to  vegetation.  The  egg  is  ^>5-%o  inch  (1.1-1.2  mm.),  the  outer  en 
velope  indistinct,  %-H  inch  (4-8  mm.),  merging  in  the  jelly  mass,  the  inne 


TREE  FROGS:  HYLIDAE  347 

envelope  Me-M.2  inch  (1.4-2.0  mm.).  The  eggs  hatch  in  4  to  5  days.  The  tad- 
pole is  medium  up  to  2  inches  (50  mm.),  tail  long,  scarlet  or  orange  vermilion 
with  black  blotches  around  the  edges  of  the  crests,  and  with  a  long  tip.  The 
tooth  ridges  are  %.  After  45  to  65  days  the  tadpoles  transform  from  June  27 
to  August  at  l/2-%  inch  (13-20  mm.). 

Journal  notes:  June  19,  1907,  Ithaca,  N.Y.  Ever  since  Sunday,  June  16,  the 
tree  toads  have  been  in  chorus.  Every  day  this  week  we  have  heard  occasional 
tree  toads  and  at  night  they  were  deafening  when  near  their  ponds.  After 
each  thundershower  they  liven  up  perceptibly.  This  evening  at  8:45  P.M.,  I 
reached  the  Veterinary  College  pond.  Frogs  were  in  chorus.  In  15  minutes  I 
had  captured  20  individuals  (including  a  mated  pair).  Found  them  also  in 
grass  near  by,  migrating  to  the  pond,  one  in  the  road  just  west.  Toads  were 
singing  here  also.  Went  out  to  cross  roads.  Here  they  were  just  as  common. 
The  log  in  southwest  corner  had  tour  perched  on  it.  To  show  how  tame  and 
how  dazed  they  are  by  the  light,  I  stroked  a  croaking  male  with  the  lighted 
end  of  my  flashlight  91  times  without  his  stirring.  He  croaked  just  the  same. 
I  could  have  repeated  the  operation.  In  a  tree  on  the  north  edge  of  the  pond 
were  four  males,  two  on  one  limb  facing  each  other. 

June  2i,  Cross  Roads  Pond,  n  A.M.  All  around  the  pond  were  tree  toad  eggs. 
I  staked  out  about  six  or  seven  areas  of  them.  The  packets  or  single  emissions 
of  eggs  maybe  6-12  inches  apart  or  only  an  inch  or  less.  Sometimes  from 
groups  of  eggs  at  more  or  less  definite  intervals  one  can  determine  the  path 
of  the  pair.  The  eggs  are  almost  invariably  at  the  surface.  Those  beneath  I  be- 
lieve were  laid  before  some  of  our  rainstorms  raised  the  water  in  the  pond. 
They  may  be  attached  to  grass  leaves,  plant  leaves,  or  sometimes  algae.  The 
packets  are  from  15-35  eggs.  Some  of  the  first  tree  toad  eggs  laid  here  are 
now  hatching.  The  wood  frog  tadpoles  are  beginning  to  develop  good-sized 
legs. 

June  i  3,  Cross  Roads  Pond.  Today  at  1 130  found  several  fresh  packets  of 
Hyla  versicolor  eggs,  attached  to  grass  at  edge  or  to  Potamogeton  leaves  in 
mid-pond.  Four  to  ten  eggs  in  a  packet.  They  were  all  at  the  surface.  Water 
surface  74°,  water  bottom  72°. 

Authorities9  corner: 

G.  H.  Loskicl,  Gen.,  1794,  p.  891.  G.  M.  Allen,  N.H.,  1899,  p.  72. 

M.  H.  Hmckley,  Mass.,  1880,  pp.  104-      F.  Overton,  N.Y.,  1914,  p.  32. 

107.  H.  Sweetman,  Mass.,  1944,  pp.  499-501, 

J.  C.  Gcikie,  Gen.,  i882(?),  pp.  217-218.      R.  L.  Hoffman,  Va.,  1946,  pp.  141-142. 

Cope's  Tree  Frog,  Western  Tree  Frog,  Chameleon  Tree  Frog 

Hyla  versicolor  chrysoscelis  (Cope).  Plate  LXXIV;  Map  20. 

Range:  Southern  Arkansas  to  east-central  and  southern  Texas. 

'The  two  forms  of  Hyla  versicolor,  the  pustulous  (typical  versicolor)  and 


348 


HANDBOOK  OF  FROGS  AND  TOADS 


Plate  LXXIV.  Hyla  vcrsicolor  chrysosce- 
lis.  i.  Female  (XO-  2,3.  Males  (Xi%). 


Plate  LXXV.  Hyla  wrightorum  (X%). 
1,3.  Male  with  throat  partly  inflated  2.  Fe- 
male in  water.  4.  Male  in  water. 


TREE  FROGS:  HYLIDAE 


349 


the  smooth  (variety  chrysoscelis)  are  distributed  over  most  of  eastern  Texas, 
but  the  present  form  seems  to  be  confined  principally  to  the  extreme  north- 
eastern counties  and  the  Trinity  and  Brazos  valleys"  (J.  K.  Strecker,  Jr.,  and 
W.  J.  Williams,  Tex.,  1928,  p.  11). 

Habitat:  Wooded  stretches  along  creeks  and  rivers. 

"In  Bowie  County  [Texas],  under  slabs  of  wood  lying  along  the  margins 
of  sloughs,  Mr.  Williams  found  three  specimens  of  chrysoscelis"  (J.  K. 
Strecker,  Jr.,  and  W.  J.  Williams,  Tex.,  1928,  p.  u). 

Size:  Adults,  i%-i%  inches.  Males,  36-43  mm.  Females,  35-49  mm. 

General  appearance:  These  frogs  are  a  smooth-skinned  version  of  our  com- 
mon tree  toad,  Hyla  v.  verstcolor.  Of  a  pair  in  hand,  each  has  a  light  spot 
below  the  eye,  legs  barred,  and  the  irregular  cross  on  the  back.  The  male  at 
present  is  a  light  grayish  tan  with  the  pattern  in  green  outlined  with  black. 
The  female  is  gray  with  the  pattern  in  dark  olive  and  black.  Both  have  the 
characteristic  orange  on  the  groin  and  concealed  portions  of  the  legs.  In  these 
two  there  is  no  black  reticulation  on  the  orange  rear  of  the  femur.  Often  the 
rear  of  the  femur  is  marked  with  very  fine  spots.  The  dark  mottling  on  the 
sides  is  very  distinct.  The  pattern  of  the  back  is  rather  less  massive  than  in  the 
average  Hyla  verstcolor.  The  male  seems  a  little  smaller  than  the  average 
Hyla  verstcolor  male.  Sometimes  the  backs  are  green. 

Color:  Male.  Becville,  Tex.,  March  27,  1925.  Orange  or  cadmium  orange  on 
side  of  groin  and  front  of  thigh,  merging  into  mars  yellow  or  xanthine  orange 
on  rear  of  thighs  and  on  the  tibia.  Spots  on  back,  calla  green.  Background 
kildare  green  or  mignonette  green. 

Another  male.  Mud  Creek,  9  miles  north  of  San  Antonio,  Tex.,  taken  by 
R.  D.  Quillm  and  A.  H.  Wright,  June  24, 1930.  Dorsum  along  sides  pale  olive- 
gray  or  pale  smoke  gray  becoming  on  back  olive-buff  or  seafoam  green  near 
edges  of  large  dorsal  spot.  Dorsal  spot  in  center  half  brown  becoming  near 
edge  dark  greenish  olive  or  dark  ivy  green.  One  forward  lateral  wing  of  spot 
dark  ivy  green.  Another  oblique  wing  from  rear  of  spot  to  each  groin,  dark 
ivy  green.  Rear  and  front  of  femur  zanthme  orange.  As  front  of  femur 
joins  body  color  it  becomes  cadmium  yellow  or  light  cadmium,  and  this  ex- 
tends along  groin.  Axil  with  touch  of  this  color  or  lemon  yellow.  Rear  of 
femur  spotted  with  finely  sized  lemon  yellow  spots.  A  few  or  a  line  of  these 
along  rear  tibial  edge.  Area  around  vent  and  along  rear  of  femur  a  short  dis- 
tance silvery  white  and  black.  Two  bars  like  dorsal  spot  across  antebrachium, 
one  across  base  of  third  and  fourth  fingers,  one  faint  one  across  brachium. 
Two  such  prominent  spots  on  femur,  two  on  tibia,  one  on  tarsus,  and  four  or 
five  on  foot.  In  front  of  vent  from  first  femur  bar  to  other  first  femur  bar  sea- 
foam  green.  Underside  of  fcmui,  hind  foot,  some  of  belly  russet-vinaceous  or 
light  russet-vinaceous.  Black  bar  from  nostril  to  eye.  Dark  area  from  eye 
through  tympanum.  Then  an  imperfect  row  of  large  dark  spots  to  groin  with 
smaller  spots  below  it.  Spot  below  eye  white,  ivory  yellow,  or  seafoam  green. 


35o  HANDBOOK  OF  FROGS  AND  TOADS 

Iris  rim  deep  seafoam  green;  iris  mainly  spotted  vmaceous-russet  with  some 
chocolate  or  burnt  umber  reticulations. 

Female.  Same  locality.  Dorsum  smoke  gray.  Dorsal  spots  and  wings  cepha- 
lic and  caudal  separate,  each  ivy  green  or  lincoln  green.  Black  stripe  from 
snout  to  eye.  Black  villa  from  eye  around  tympanum  halfway  to  groin.  Below 
this  a  light  drab  band  from  tympanum  almost  to  groin.  Black  line  above  in 
groin  breaks  into  black  spots  with  light  cadmium  below  it.  Spot  below  eye 
white.  Other  colors  more  or  less  as  in  male.  Throat,  belly,  and  underside  of 
fore  limbs  white. 

Structure:  Cope's  original  description  (Hyla  jemoralts  chrysoscehs)  is  as 
follows:  "Hyla  femorahs  Daudin.  A  specimen  larger  than  the  largest  indi- 
viduals I  have  previously  seen;  differs  also  in  the  greater  extent  of  the  palma- 
tion  of  the  fingers,  and  in  the  coloration  of  the  concealed  surface  of  the  femur. 
In  eastern  specimens  the  posterior  face  of  the  femur  is  brown,  with  rather 
small  yellow  spots;  in  this  form  it  is  yellow  with  a  blackish  coarse  reticulation, 
which  only  extends  to  the  lower  surface  of  the  proximal  half  of  the  thigh.  The 
sides  have  a  double  row  of  small  black  spots,  which  enclose  a  yellow  band. 
This  is  probably  a  subspecies  and  may  be  distinguished  by  the  name  of 
chrysoscehs.  One  specimen  as  large  as  a  large  Hyla  verstcolor  was  taken  by 
Mr.  Boll  near  Dallas"  (E.  D.  Cope,  Tex.,  1880,  p.  29). 

"Five  specimens  from  Helotes  were  in  the  Marnock  collection.  These  speci- 
mens are  perfectly  smooth  above  and  resemble  in  every  particular  typical 
specimens  of  chrysoscehs  from  the  type  locality.  This  is  the  common  Hyla  of 
middle  and  northern  Texas"  (J.  K.  Streckcr,  Jr.,  Tex.,  1922,  p.  15). 

Voice:  It  is  a  loud  resonant  trill. 

"This  species  is  fairly  abundant  near  Houston  and  I  have  the  following 
notes  on  it:  February  13, 1918.  Have  heard  several  in  the  past  few  days  calling 
from  trees  in  camp,  but  have  not  seen  any  yet.  April  15.  Collected  one  speci- 
men, a  male  that  was  calling  from  the  branch  of  a  pine  tree.  April  24.  Found 
one  calling  in  rain  pool  where  H.  sqturella  was  breeding.  I  heard  them  fre- 
quently on  warm  evenings,  answering  each  other  from  trees  in  the  woods 
near  camp.  After  the  first  of  May  they  became  silent  and  I  heard  and  saw  no 
more  of  them  for  the  season.  If  they  laid  eggs  in  any  of  the  pools  near  camp 
they  made  no  such  noise  about  it  as  they  do  in  the  North"  (P.  H.  Pope,  Tex., 

i9'9>  P-  95)- 

"2/17-  Heard  Hyla  v.  chrysoscehs  calling  tonight— evening,  ist  this  spring. 

"Feb.  18.  Cold.  No  frogs  or  toads. 

"Specimen  1948.  H.  v.  chrysoscehs.  7/15/44  from  damp  ground  under  wa- 
ter spigot  at  house  3"  deep. 

"Somerset,  Tex.  y1/:*  mi.  S.W.  1944.  Sept.  2. 

"Specimen  1951-1956  inc.  Hyla  v.  chrysoscehs. 

"I  have  been  hearing  a  strange  frog  or  (toad)  note  since  Aug.  27,  from  the 
Geo.  Foster  tank  or  pond  about  %  mi.  S.E.  of  our  house.  There  had  been 


TREE  FROGS:  HYLIDAE  351 

much  rain  from  Aug.  22  to  Aug.  28,  clear  since  8/28.  Tonight  I  went  to  the 
tank  and  heard  a  few  of  these  calls.  After  searching  I  found  them  to  come 
from  trees  and  bushes  and  a  tree  fallen  in  the  water.  I  caught  two  that  I 
thought  were  the  croakers.  Hylas,  somewhat  small.  Finally  I  was  watching 
two  Hylas,  one  was  croaking  the  other  about  6"  away.  Female  was  silent,  sud- 
denly the  croaking  male  began  calling  the  strange  notes:  uttered  fully  twice  as 
quick  as  the  ordinary  notes  and  an  altogether  different  tone,  clear,  shriller, 
2  to  4  notes;  ('qui  qui  qui')  sometimes  there  was  a  low  gurgling  preceding 
the  call  notes;  most  times  the  notes  were  low  but  now  and  then  they  could 
be  heard  a  long  distance,  clearly  at  our  house,  %  mile  away.  This  satisfied  me 
that  the  strange  notes  were  made  by  the  male  H.  v.  chrysoscelis  during  mating 
times.  All  the  males  (6)  heard  that  I  saw  were  near  females.  On  a  horizontal 
limb  ($"  diameter)  6  feet  up  were  5  hylas,  2  females,  }  males.  They  were  close 
together,  all  five  within  10  inches.  One  male  uttered  these  calls.  It  was  along- 
side of  a  female,  'nudged*  the  female  a  few  times.  This  pair  collected,  as  was 
the  pair  first  watched  and  2  other  single  females  (?)  caught  near  where  these 
calls  heard.  I  have  heard  these  strange  notes  a  few  times  before"  (notes,  A.  J. 
Kirn,  Somerset,  Tex.,  1944). 

Breeding:  They  breed  from  middle  March  to  July.  The  tadpoles  transform 
at  %  inch  (16  mm.). 

"This  spring  I  found  this  tree-toad  breeding  in  small  rock-bound  pools  in 
a  gravel  pit.  The  tadpoles  were  light  yellow.  Specimens  collected  April  21  st 
had  the  hindhmbs  well  developed"  (J.  K.  Strecker,  Jr.,  Tex.,  19103,  p.  118). 

Journal  notes:  March  24,  1925,  Beevillc,  Tex.  The  air  is  resounding  with 
Psetidacris.  We  heard  a  few  scattering  Hyla  versicolor  along  the  roadside 
ditches  or  in  the  distance  and  captured  three  males,  but  later  lost  one.  They 
are  along  these  roadside  ditches  but  not  in  them.  They  look  quite  green  at 
night.  We  captured  one  on  the  bole  of  a  mesquite  near  the  pond.  Meadow 
frogs  are  croaking  loudly  in  these  ditches  and  narrow-mouthed  toads  arc 
common. 

June  14,  1950,  Becville,  Tex.,  at  night.  We  were  about  to  leave  the  pond 
when  we  heard  a  H.  versicolor.  We  started  for  it  when  we  heard  two  more 
at  a  distance.  One  we  caught.  It  has  spotting  on  the  rear  of  the  femur,  very 
fine  spots  down  the  outer  half  of  the  femur.  When  we  reached  the  house  we 
lost  the  H.  versicolor. 

June  22.  We  went  out  with  R.  D.  Quillin.  At  Mud  Creek,  9  miles  north  of 
San  Antonio,  we  heard  many  Microhyla  olivacea,  Rana  piptens,  a  few  Bufo 
valliceps,  and  several  Hyla  versicolor  chrysoscelis.  We  sought  the  tree  toads. 
The  first  one  was  too  high  in  the  oaks.  The  next  one  we  picked  up  on  mud 
near  the  water.  Then  we  heard  a  croaker  near  the  spot  but  the  one  on  the  mud 
proved  a  ripe  female.  In  another  oak  tree  we  found  a  male,  3  feet  up.  This  we 
put  with  the  female.  When  we  returned  to  the  house  they  were  mated  axillary 
fashion. 


352  HANDBOOK  OF  FROGS  AND  TOADS 

June  26,  Waco,  Tex.  I  saw  some  of  Strecker's  H.  v.  chrysoscelis.  He  pro- 
nounced my  San  Antonio  live  Hyla  versicolor,  H.  v.  chrysoscelis.  These  are 
smooth  and  quite  greenish.  In  three  H.  v.  chrysoscelis  in  the  Baylor  collection 
the  suborbital  spot  is  obscure  or  absent  or  present  on  one  side  and  absent  on 
the  other.  The  femoral  reticulation  is  more  pronounced  than  in  my  H.  v. 
chrysoscelis  from  San  Antonio. 

Authorities'  corner: 
J.  K.  Strecker,  Jr.,  Tex.,  19103,  pp.  117- 

118. 

"In  the  southern  and  western  specimens  there  is  a  tendency  to  the  replacing 
of  the  brown  reticulation  on  the  yellow  ground  of  the  posterior  face  of  the 
thighs  by  a  number  ot  subcircular  golden  spots  in  the  brown  ground,  as  in 
the  Hyla  jcmoralts,  although  northern  specimens  sometimes  show  traces  of 
it.  This  is  very  evident  in  specimens  from  Prairie  Mer  Rouge  and  Tangipahoa 
River,  Louisiana,  and  Dallas,  Texas.  As  a  general  rule,  too,  the  portions  of 
the  limbs  concealed  or  in  contact  with  each  other  when  flexed,  are  in  northern 
specimens  more  fully  marbled  with  yellow,  even  covering  the  whole  inner 
surface  of  the  tibia  and  the  light  interspaces  more  or  less  angular,  while  in  the 
Stntlisca  bandmii  and  the  southern  and  western  specimens  of  H.  versicolor 
the  amount  of  marbling  is  less,  and  the  interspaces  are  often  reduced  to  small 
circular  spots.  I  have,  however,  been  unable  to  characterize  them  as  more  than 
a  variety,  to  which  I  give  the  name  of  H.  v.  chrysoscelis''  (E.  D.  Cope,  Gen., 
1889,  pp.  374-375). 

"It  is  easy  to  be  confused  in  trying  to  solve  Cope's  original  Hyla  versicolor 
chrysoscelis,  H.  v.  versicolor,  and  Hvla  v.  phacocrypta.  Viosca  positively  as- 
serts that  H.  v.  phaeocrypta  type  is  H.  v.  versicolor.  Stejneger  early  pointed 
out  that  H.  v.  chrvsoscelis  could  not  be  H.  fcmoralis. 

"When  after  H.  avivoca  with  Viosca  and  Chase  on  June  n,  1930  in  Tick- 
faw  and  Tangipahoa  Country,  'we  took  in  road  ditches  several  H.  v.  versi- 
color. Are  they  H.  v.  chrysoscelis?  Quite  smooth.  The  female  is  very  large  in 
contrast  to  the  males  captured.  Started  for  H.  avivoca  nearby.  It  is  not  the 
H.  versicolor  of  Louisiana,  but  is  quite  different  and  different  in  ecological 
habits.  Yet  if  I  read  Cope's  recharacterization  (1889)  of  H.  v.  chrysoscelis 
(1880)  (partially  from  Louisiana,  Tangipahoa  River — this  very  country  I  am 
in)  one  might  suspect  Cope  might  have  had  some  H.  avivoca.  If  so  his  Tangi- 
pahoa River,  La.,  and  Dallas,  Texas,  description  is  of  a  composite  form.  Cope 
(1880)  put  his  form  in  H.  femoralis,  but  Hyla  avivoca  never  was  H.  femoralis 
in  habit.  It  is  found  in  tupelo  gums,  H.  femoralis  in  pine  woods.  Each  of  these 
and  H.  versicolor  are  in  this  region  of  Louisiana.  .  .  .'  Strecker  redescribes 
H.  v.  chrysoscelis.  He  calls  it  a  smooth-skinned  frog  with  a  yellow  suborbital 
spot.  He  also  alludes  to  Cope's  character  of  fine  femoral  spots.  I  have  taken 
H.  versicolor  from  Beeville  to  north  of  San  Antonio.  I  believe  that  it  is  the 
form  Strecker  has  interpreted  as  H.  v.  chrysoscelis.  I  showed  him  some  live 


TREE  FROGS:  HYLIDAE 


353 


specimens  from  Mud  Creek  9  miles  north  of  San  Antonio,  Texas.  These 
seemed  much  like  Waco,  Texas,  specimens  he  showed  me.  Both  lots  (mine 
and  his)  are  smooth-skinned  but  some  have  no  suborbital  spot,  or  occasionally 
they  have  a  line  from  nostril  to  eye  and  backward.  The  Louisiana  material 
needs  to  be  restudied.  At  present  we  will  hold  to  the  Dallas  (Boll)  material  as 
the  center  of  its  (H.  versicolor  chrysoscehs)  eastern  Texas  to  Arkansas  range" 
(A.  H.Wright,  MS,  1930). 

Recently  H.  M.  Smith  and  Bryce  C.  Brown  assigned  the  Hyla  versicolor  of 
the  "Lower  Mississippi  Valley  west  through  eastern  Texas  to  about  Leon  and 
Austin  counties"  to  H.  versicolor  chrysoscelis.  This  they  diagnose  as  follows: 
"Like  H.  v.  versicolor  except  rear  surface  of  thigh  more  extensively  dark 
marked,  leaving  only  isolated,  circular  light  areas  of  moderate  size."  This 
seemingly  is  in  agreement  with  Cope's  saying  he  had  it  from  Dallas,  Tex., 
and  Tangipahoa  River,  La.  The  new  form,  H.  versicolor  sandersi,  after  our 
friends  Mr.  and  Mrs.  Ottys  Sanders,  they  restrict  as  follows:  "The  Balcones 
escarpment  and  its  vicinity  in  central  Texas,  south  at  least  to  Atascosa  County, 
east  to  Bastrop  County,  north  to  Travis  County  and  no  doubt  to  McLellan 
County."  "The  Texan  subspecies  may  be  diagnosed  as  follows :  rear  of  thigh 
light  (orange  in  life),  with  fine  white  flecks,  entirely  lacking  dark  marks  ex- 
cept at  extreme  medial  border;  fingers  not  or  only  barely  perceptibly  webbed; 
skin  smooth;  maximum  snout-vent  length  lesser,  4?  mm.  in  males,  48  mm.  in 
females.  .  .  ." 

Dusky  Tree  Toad 

Hyla  versicolor  phaeocrypta  (Cope).  Plate  LXXVI. 

Range:  Specimens  from  Nashville,  Tenn.,  Olive  Branch,  111.,  Mt.  Carmel, 
111.,  Olney,  111.,  Gull  Lake,  Brainerd,  Minn.,  and  Springfield,  S.D.,  have  been 
assigned  to  this  form. 

Habitat:  River  valley  and  lake  shore. 

Size:  Adults,  i%-i%  inches.  Males,  as  large  as  33  mm.  Females,  as  large 
as  36  mm. 

General  appearance:  This  is  another  of  Cope's  H.  versicolor-H.  femoralis 
puzzles  based  on  preserved  material.  It  is  as  yet  unsolved;  however,  it  is 
probably  not  a  good  form.  Cope's  original  description  follows : 

"A  single  specimen  of  a  strongly  marked  variety  of  this  species  was  sent 
to  the  National  Museum  from  Mt.  Carmel,  111.,  by  Lucien  M.  Turner  (No. 
12074).  It  is  smaller,  having  the  average  dimensions  of  H.  femoralis.  The  color 
is  a  dark  brown,  with  three  rows  of  large  approximated  darker  brown  spots. 
The  groin  and  concealed  faces  of  the  thigh  are  yellowish  brown,  with  a  very 
scanty  speckling  of  darker  brown,  very  different  from  the  usual  coarse  netted 
pattern.  At  first  sight  one  suspects  this  to  be  a  specimen  of  Hyla  femoralis, 
but  it  possesses  all  the  essential  characters  of  the  integument  and  feet  of  the 


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TREE  FROGS:  HYLIDAE  355 

H.  versicolor,  as  pointed  out  in  the  analytical  table  of  the  genus,  including 
also  the  light  spot  under  the  eye.  It  may  be  called  H.  v.  phaeocrypta9  (Cope, 
Gen.,  1889,  p.  375). 

In  hurried  visits  to  three  of  its  localities,  we  have  met  questionable  forms. 
Live  material  from  Wabash  Valley  is  our  greatest  need.  Viosca  now  pro- 
nounces the  type  of  Hyla  phaeocrypta  a  poorly  preserved  Hyla  versicolor. 
What  is  this  form?  In  the  last  fifteen  years  little  has  appeared  to  add  to  the 
solution. 

From  Itasca  Park,  Minn.  This  is  a  smooth-skinned  tree  toad,  gray  with 
prominent  crossbars  on  legs  and  varying  types  of  spots  on  the  back.  In  some 
these  spots  assume  a  cruciform  pattern  or  are  arranged  in  three  or  four  rows. 
There  is  a  white  or  ohvine  spot  below  the  eye.  From  the  eye  backward  ex- 
tends an  oblique  band  of  gray  or  vmaceous  bordered  above  with  black.  There 
is  much  cadmium  orange  or  cadmium  yellow  on  rear  and  front  of  femur,  on 
underside  of  foot,  in  groin,  and  in  axil.  From  Walhalla,  N.D.  In  darkness 
and  moisture,  this  frog  was  dark  grayish  olive  and  very  little  dorsal  pattern 
showed.  Brought  into  the  light,  he  is  rapidly  becoming  lighter  and  the  dorsal 
pattern  is  coming  out.  He  lacks  the  cruciform  patch  of  typical  versicolor. 
There  are  about  four  rows  of  irregular  spots  on  the  back. 

These  and  the  Kansas,  South  Dakota,  and  Wisconsin  records  are  likely  to 
prove  variant  H.  v.  versicolor. 

Structure:  For  those  who  wish  to  follow  the  relative  measurements  of  Hyla 
avtvoca,  Hyla  v.  versicolor,  and  Hyla  v.  phaeocrypta  (Illinois  northwest- 
ward), the  following  table  ot  1930  is  given: 

H.a.  H.p.  H.v.  H.a.  H.p.  H.v. 

No.  USNM  75026  70644  C.U.  75018  70643  3636 

Sex  Male  Female'5  Female  Female 

Length  28  28  28  36  36  36 

Head-tympanum  10             9  10  n  12  12.5 

Head-angle  of  mouth  9.5          7             9  10  10  11.5 

Width  head  n  10  12  12  13.5  14 

Snout  5            4.5          5-5  5-5  6  6 

Eye  334  4-5  3-5  4 

Interorbital  space  3.5          3.5          2.5  5  4  5 

Upper  eyelid  3.5          2.5          2.5  4  3.5  4 

Tympanum  2233  2.5  2.5 

Intertympanic  space  10.5          8.5          9  n  11.5  11.5 

Internasal  space  3.5          333-53  4 

Fore  limb  18  14  20  20  17  24 

ist  finger  4.5          4            3-5  5-5  4-5  6 

2nd  finger  6            4.5          5  7.5  5  8-5 

3rd  finger  9.5          6            8  10  7  n 

4th  finger  7.5          5             6  8.5  6  9 

Hind  limb  42  35  44  55  53  54 


356  HANDBOOK  OF  FROGS  AND  TOADS 

H.a.  H.p.  H.v.  H.a.  H.p.  H.v. 

Tibia  15          15  16  18.5  19 

Foot  with  tarsus  19.5        16  24.5  24  27 

Foot  without  tarsus  14  9  14  14.5  14  15.5 

isttoe  4-5435  6  7 

2nd  toe  7  6  4.5  8  8  10 

3rd  toe  10.5         8  8  12.5  10  12 

4th  toe  14  9  ii  14.5  12  15.5 

5th  toe  ii  8  8  12  9.5  12 

Remarks:  The  material  which  comes  into  this  consideration  is: 
USNM  no.  12074,  acl-  Mt-  Carmel,  111.  L.  M.  Turner  (type). 
USNM  no.  70642, 34  mm.  3  Gull  Lake,  10  mi.  n.  Brainerd,  Minn. 
USNM  no.  70643, 36  mm.  $  Gull  Lake,  10  mi.  n.  Brainerd,  F.  H.  Uhler. 
USNM  no.  70644, 28  mm.  yg.  Gull  Lake,  10  mi.  n.  Brainerd,  F.  H.  Uhler. 
USNM  no,  71588, 10  mi.  ne.  Olney,  Richland,  111.,  Robt.  Ridgway. 
USNM  no.  68719,  28  mm.  yg.  or  ?  Springfield,  S.D.,  E.  C.  O'Roke. 

Mittleman  kindly  showed  us  his  manuscript.  It  is  a  good  paper  and  he  may 
be  right.  In  the  preceding  table  we  compared  one  28-mm.  and  one  36-mm. 
specimen  of  each  form,  six  examples  in  all.  We  still  retain  H.  v.  phaeocrypta 
in  this  book  for  someone  to  catch  the  Wabash  frog  in  the  act  and  then  for  the 
same  person  the  same  season  to  familiarize  himself  with  the  song  of  H.  avi- 
voca. 

Some  of  Mittleman's  statements  and  our  comments  follow: 

1.  "In  addition  I  have  heard  the  call  of  this  species  (H.  phaeocrypta,  i.e., 
avivoca)  in  the  gum  trees  of  the  mud  flats  along  the  Wabash  River  on  the 
outskirts  of  Terre  Haute,  Indiana"  (p.  34).  Quite  likely  he  did  hear  it.  Why 
compare  this  memory  with  an  old  preserved  specimen  ? 

2.  "The  highly  distinctive  voice  of  phaeocrypta  (avivoca)  need  be  confused 
with  no  other  North  Ameiican  hylid  save  possibly  H.  cruet  fer"  (p.  35),  Too 
comprehensive  a  statement.  Authors  speak  of  "breeding  song,"  "rain  song," 
"chorus  note,"  and  other  notes  for  one  species.  One  or  two  species  may  vary 
an  octave, 

3.  "Phaeocrypta.  Call,  a  piping,  bird  whistle.  Versicolor.  Call,  a  toad-like 
trill"  (p.  36).  To  DeKay  versicolor  had  a  ventnloqual  quality,  to  Cope  a  reso 
nant  trill,  to  Fowler  a  jerking  cry,  to  Dickerson  a  cat  purr  or  lamb  bleat,  tc 
Overton  a  musical  trill  like  a  low-pitched  whistle  (trill  and  whistle  combined) 
and  a  turkey  call,  to  Harper  (1921)  a  vigorous  high-pitched  trill,  and  so  on 
Did  Turner  actually  catch  a  birdlike  Hyla  at  Mt.  Carmel?  What  is  birdlike— 
Overton's  turkey  root,  Viosca's  and  Wright's  woodpecker  call  for  avivoca,  o\ 
somebody's  musical  thrushlike  notes ?  What  birds  do  we  choose  ? 

4.  "If  Harper's  specimen  is  truly  referable  to  phaeocrypta  it  probably  indi 
cates  a  significant  trend  in  Atlantal  coastal  populations.  It  seems  much  mor< 
likely  that  it  is  simply  a  versicolor"  (p.  34).  Harper  at  Louisville,  Ga.,  took  , 


TREE  FROGS:  HYLIDAE  357 

female  avivoca  of  49  mm.,  5  mm.  longer  than  what  we  reported  in  1933,  or 
6  or  7  mm.  more  than  Mittleman's  specimens.  Why  not?  We  have  been  in 
the  field  many  times  with  the  four  veterans  who  know  H.  avivoca  best, 
P.  Viosca,  Jr.,  O.  C.  Van  Hyning,  A.  F.  Carr,  Jr.,  and  F.  Harper.  They  know 
the  southeast  frogs  alive,  in  alcohol,  and  theoretically. 

Color:  Male.  Itasca  Park,  Minn.,  Aug.  26,  1950.  The  spots  of  back  are  deep 
olive-gray  with  dark  margins.  These  spots  consist  of  four  irregular  dorsal 
rows,  and  a  bar  from  each  eyelid  extending  obliquely  backward  and  inward 
but  not  meeting  its  fellow.  The  background  of  the  back  is  smoke  gray  with 
black  line  from  snout  to  eye,  resumed  back  of  eye,  and  extending  over  tym- 
panum and  arm  insertion  and  along  side.  The  area  below  it  and  involving  the 
tympanum  is  drab-gray  or  light  purplish  gray.  This  area  on  its  lower  portion 
is  separated  from  the  light  cadmium  of  the  axil  by  oblique  black  border.  The 
spot  below  the  eye  is  white.  Transverse  area  above  vent  is  white,  bordered 
below  by  black.  Below  this  is  a  pale  flesh  area.  Rear  of  femur  has  prominent 
cadmium  orange  spots  with  amber  brown  interspaces.  Underside  of  tibia  is 
amber  brown  with  a  few  orange  spots.  So  also  is  the  inside  of  tibia  and  foot. 
The  groin  is  mainly  cadmium  yellow.  The  eye  is  spotted  with  congo  pink. 
The  throat  is  light  grayish  olive  with  rest  of  venter  white. 

Another  male.  Walhalla,  N.D.  Sept.  i,  1930.  The  general  dorsal  color  is  dark 
olive-gray,  becoming  olivine  on  top  of  foreleg,  on  top  of  femur  and  tibia,  and 
on  sides  above  the  dark  edge  of  lateral  band.  The  spot  below  the  eye  is  of  the 
same  color.  Band  from  eve  through  tympanum  and  over  arm  insertion  is  light 
drab.  The  tympanum  is  grayish  olive.  This  lateral  band  is  bordered  by  black. 
Just  above  groin  is  a  prominent  black-edged  spot;  its  center  is  olivine.  There 
is  a  slight  touch  of  clear  orange  or  cadmium  yellow  in  axil,  groin,  and  front  of 
femur.  The  rear  of  femur  is  finely  spotted  with  cadmium  yellow,  on  a  xan- 
thme  orange  or  amber  brown  background;  same  on  underside  of  tibia  and 
inner  side  of  foot.  The  femur  is  slightly  barred,  the  tibia  more  so.  There  is  a 
V  from  one  eyelid  to  the  other.  The  spots  of  back  are  black  or  with  citrine  cen- 
ters. The  underside  of  lower  jaw  is  olivine  with  some  dark  clashes.  The  lower 
throat  is  deep  grayish  olive.  The  rest  of  under  parts  are  white  except  for  im- 
dersurfaces  of  hind  limb,  which  arc  pale  flesh  color.  Just  above  anus  is  a  light 
lumiere  green  area  bordered  below  by  black,  and  this  in  turn  by  pale  grayish 
vinaceous,  below  that  a  silvery  pebbling,  some  of  which  appears  also  on  tarsus. 
The  iris  is  black,  heavily  spotted  with  vinaccous-tawny;  pupil  rim  broken 
above,  below,  behind,  and  in  front;  pupil  rim  is  light  greenish  yellow  above. 

Female.  Itasca  Park,  Aug.  26,  1930.  The  spots  on  back  are  light  olive-gray  or 
tea  green  outlined  with  dotted  lines  of  black.  The  bars  on  femur,  tibia,  and 
antebrachium  are  the  same  color.  The  general  background  of  back  is  pale 
drab-gray.  From  the  eye  there  extends  backward  along  the  side,  an  ecru-drab 
or  pale  brownish  drab  band  bordered  above  by  black-dotted  line.  The  spot 
below  the  eye  is  white  or  marguerite  yellow.  The  tympanum  is  light  cinna- 


358  HANDBOOK  OF  FROGS  AND  TOADS 

mon-drab.  The  front  of  femur,  underside  of  tibia,  and  rear  lower  half  of  femur 
is  cadmium  yellow  with  same  color  in  groin  and  to  slight  extent  in  axil.  This 
cadmium  yellow  forms  fine  spots  on  rear  of  femur  and  slightly  on  rear  and 
forward  edge  of  tibia,  with  interstices  of  sudan  brown  or  antique  brown.  The 
iris  is  heavily  spotted  with  japan  rose.  Ventral  surfaces  are  white.  With  this 
frog,  the  spots  on  the  back  were  a  large  irregularly  cruciform  one  on  the  for- 
ward part  of  the  back  and  two  smaller  ones  to  the  rear  and  laterad. 

(These  three  specimens,  two  from  Itasca  Park,  Minn.,  and  one  from  Wal- 
halla,  N.D.,  may  eventually,  of  course,  be  referred  to  H.  v.  versicolor.) 

Voice:  "As  it  is  impossible  to  revive  the  voice  of  Copy's  type  specimen,  some 
one  must  hear  and  collect  its  descendants  in  or  near  the  type  locality.  Fortu- 
nately, this  has  virtually  been  done  through  the  kindness  of  Mr.  Karl  P. 
Schmidt,  of  the  Field  Museum,  who  has  called  my  attention  to  a  series  of 
specimens  from  Olive  Branch,  Illinois,  in  that  museum,  and  these'  seem  to  be 
the  same  as  Cope's  phaeocrypta.  The  field  notes  of  Mr.  C.  M.  Barber  which 
were  submitted  by  Mr.  Schmidt  are  especially  interesting,  as  they  may  supply 
the  missing  evidence: 

"  Whilst  collecting  Hyla  cincrea  with  a  light  in  a  boat,  on  Horseshoe  Lake, 
May  23,  1907  with  Dr.  Chatham,  my  attention  was  called  to  a  bird-like  cry 
coming  from  trees  and  bushes  overhead.  At  first  I  thought  it  resembled  the 
rattle  of  Acns,  but  it  has  a  bird-like  quality  quite  peculiar.  Several  searches 
were  unsuccessful,  but  finally  we  were  able  to  find  the  sober-colored  little 
frog  when  he  uttered  his  cry,  fearlessly,  within  a  few  feet,  and  with  the  light 
fully  on  him.  Their  note  is  strong  and  penetrating  and  more  pleasing  than 
any  Hyla  that  I  know.  I  had  previously  taken  one  in  the  grass  near  the  lake, 
but  mistook  it  for  a  juvenile  versicolor  "  (P.  Viosca,  Jr.,  La.,  1923^  pp.  97-98). 

In  1924  Robert  Ridgway  (111.,  p.  39)  writing  under  the  title,  "Additional 
Notes  on  Hyla  phaeocrypta  (D),"  reminisced  as  follows: 

"Mr.  Viosca's  observations  on  Hyla  phaeocrypta  Cope  in  Copeia  122  recall 
to  my  memory  an  experience  o£  many  years  ago  in  the  bottom-lands  of  the 
Wabash  and  tributary  streams  near  Mt.  Carmel,  111.  The  bird-like  notes  de- 
scribed by  Mr.  Viosca  were  frequently  heard  by  me  in  the  woods,  so  distinctly 
bird-like  that  I  was  constantly  looking  for  the  unknown  bird;  in  fact,  there 
was  a  "standing  reward"  offered  to  my  boy  friends  for  a  specimen  never  dis- 
covered. Some  years  later,  when  my  father  moved  to  a  farm  near  Wheatland, 
Ind.,  I  learned  from  him  that  it  was  a  tree-frog.  Whether  Hyla  phaeocrypta 
or  not,  the  species  is  so  abundant  in  the  bottom-land  woods  that  at  certain 
seasons  (if  my  memory  is  not  at  fault,  in  late  summer  or  autumn)  its  clear, 
loud  piping  notes — as  different  as  possible  from  the  croak  of  H.  versicolor — 
were  the  dominant  and  often  the  only  sound  to  be  heard"  (R.  Ridgway,  111., 
1924,  p.  39). 

Breeding:  Nothing  of  record. 

Journal  notes:  Sept.  13, 1929,  Calhoun,  111.  While  collecting  in  the  northern 


TREE  FROGS:  HYLlDAE 


359 


gopher  frog  areas  which  Lee  Ackert  showed  us,  we  came  to  a  pond.  When 
about  to  leave,  we  heard  a  Hyla  versicolor.  It  was  in  a  bush  in  the  deep  water 
of  the  pond,  and  so  distinctly  H.  v.  versicolor  that  we  passed  it  by  in  our 
search  for  R.  areolata  circulosa.  Later  we  wondered  if  it  might  not  have  been 
Ridgway's  H.  phaeocrypta. 

Sept.  14.  Went  to  Mt.  Carmel,  111.,  and  New  Harmony,  Ind.,  for  a  day  but 
heard  nothing  like  the  reputed  H.  v,  phaeocrypta. 

Aug.  26,  1930,  Gull  Lake,  Minn.  Wanted  to  take  some  of  the  queer  hylas 
F.  M.  Uhler  secured  here,  but  found  instead  almost  every  other  species  of  the 
region. 

Sept.  7, 1930.  Drove  around  the  country  at  Mt.  Carmel  and  listened  at  night 
and  by  day,  but  discovered  no  queer  Hyla  notes  such  as  those  of  H.  v.  phaeo- 
crypta  are  reputed  to  be. 

Oct.  7,  1929.  A  small  male  tree  toad  from  northern  Wisconsin  (from  Caro- 
lyn Weber),  we  provisionally  placed  in  this  group.  Our  general  notes  on  it  are: 
This  is  a  small  specimen  of  tree  toad.  It  is  brownish  gray  on  the  back  with 
irregular  black  markings,  forming  an  irregular  and  imperfect  cross.  A  dark 
band  extends  from  eye  downward  across  the  tympanum  and  along  the  side. 
The  top  of  this  band  is  marked  with  a  broken  black  line  bordered  above  by 
light.  The  sides  are  speckled  brown.  The  legs  are  barred,  the  throat  is  dark. 
There  is  orange  on  rear  ot  hind  legs,  in  groin,  and  in  axilla. 

Not  until  we  have  several  of  these  frogs  alive  from  Mt.  Carmel,  Olive 
Branch,  and  Gull  Lake,  will  we  feel  satisfied  or  competent  to  weigh  the  posi- 
tion of  this  species.  Is  it  a  separate  form  or  H.  versicolor  versicolor,  H.  v. 
chrysoscelis,  H.  avivoca,  or  Pseudacns  brachyphona? 

Authorities9  corner:  In  1923  while  provisionally  placing  his  later  H.  avivoca 
under  H.  phaeocrypta,  Viosca  wrote:  "As  virtually  all  the  distinguishing  char- 
acters of  my  new  frog  were  lost  in  the  preservative,  its  general  resemblance  to 
Hyla  versicolor  made  comparison  with  museum  material  difficult,  especially 
since  versicolor  is  such  a  variable  species.  Except  for  the  smaller  size,  which 
could  easily  have  been  due  to  age,  there  is  nothing  in  Cope's  description  of 
phaeocrypta  that  would  suggest  my  specimens.  Dr.  Stejnegcr  has  compared 
specimens  of  my  species  with  Cope's  type  of  phaeocrypta  with  the  result  that 
structurally  at  least  he  could  not  tell  them  apart,  yet  this  in  itself  was  not  con- 
clusive, as  my  specimens  also  exhibit  a  structural  resemblance  to  small  speci- 
mens of  Hyla  versicolor  chrysoscehs.  .  .  .  Hyla  phaeocrypta  .  .  .  should  how- 
ever be  rediscovered  in  the  type  locality  and  the  voice  noted.  Although  there 
are  apparent  difTerences  in  the  specimens  at  hand,  between  the  phaeocrypta 
of  Illinois  and  the  bird-voiced  Hyla  of  Southeast  Louisiana,  it  is  difficult  to 
pass  an  opinion  with  the  limited  material  available.  These  also  must  be  con- 
sidered the  same  until  both  forms  can  be  compared  in  life,  or  better,  be  heard 
by  the  same  person"  (P.  Viosca,  Jr.,  La.,  igijb,  pp.  97-98). 

In  describing  his  Louisiana  Hyla  as  Hyla  avivoca  not  Hyla  phaeocrypta 


360  HAKDfiOOK  OF  FROGS  AND  TOADS 

Viosca  said  in  1928:  "The  species  described  herein  had  been  called  tentatively 
Hyla  phaeocrypta  Cope  pending  its  comparison  by  the  writer  with  Cope's 
type  of  phaeocrypta,  as  well  as  with  other  North  American  Hyla  types  in 
the  National  Museum.  I  have  had  an  opportunity  recently  to  make  such  a 
study  in  the  National  Museum  and  found  the  Louisiana  form  to  be  a  species 
distinct  from  any  North  American  Hyla  heretofore  described.  Further,  I 
found  the  type  of  Hyla  versicolor  phaeocrypta  Cope,  U.S.N.M.  No.  12074,  to 
be  a  fairly  typical  specimen  of  Hyla  versicolor,  poorly  preserved  as  to  texture 
and  color,  but  well  within  the  range  of  individual  variations  normally  ex- 
hibited by  that  species.  The  typical  cruciform  pattern  of  versicolor,  though 
faint,  is  readily  discernible,  and  the  structural  characters  pointed  out  by  Cope 
place  it  unquestionably  with  that  species"  (P.  Viosca,  Jr.,  La.,  1928,  p.  89). 

Note  on  W.  T.  Neill's  (1948)  significant  paper  on  an  eastern  race,  Hyla 
phaeocrypta  ogechiensis  for  Georgia  and  Carolina  specimens:  Having  visited 
the  type  locality  of  Hyla  phaeocrypta  without  success,  we  want  evidence  from 
someone  with  success  in  Mt.  Cartnel  and  the  lower  W  abash  country.  Northern 
specimens  arc  doubtless  versicolor  but  avwoca-phaeocrypta  is  not  yet  proved. 
Avivoca's  range  may  extend  in  the  Upper  Coastal  area  to  North  Carolina. 

Sonora  Tree  Frog,  Wrights'  Tree  Frog,  Sonora  Hyla 
Hyla  wnghtontm  Taylor.  Plate  LXXV;  Map  iS. 

Range:  North-central  Arizona,  New  Mexico,  Chihuahua  possibly  to  Texas. 

"To  date  the  author  has  found  this  frog  only  on  the  southern  forested  edge 
of  the  Colorado  Plateau  from  the  vicinity  of  Williams  east  to  McNary.  The 
full  range  extends  cast  into  Texas  and  south  into  Mexico. 

"The  Colorado  Plateau  extends  roughly  east  and  west  through  central  Ari- 
zona into  New  Mexico.  The  altitude  is  approximately  7000  feet  above  sea  level. 
In  general  the  southern  part  is  in  the  transition  zone  and  is  covered  by  typical 
vegetation  including  a  broad  belt  of  ponderosa  pine  with  small  areas  of  Doug- 
las fir,  white  fir,  Mexican  white  pine,  and  other  irees.  .  .  .  No  specimens  of 
Hyla  wrightontm  were  found  in  the  isolated  Final  Mountains,  south  of  Globe, 
although  they  reach  fiom  upper  Sonoran  through  the  transition  zones,  rang- 
ing from  4000  to  8000  feet  in  elevation.  No  frogs  were  found  in  the  large 
springs  at  Pinetop,  whose  waters  have  a  temperature  of  about  50°  F.  The 
waters  of  open  ponds  arc  warmed  during  the  day,  and  remain  at  a  higher 
temperature. 

"The  lower  limit  of  the  distribution  of  the  species  in  this  region  seems  to 
be  defined  by  the  rim  of  the  Colorado  Plateau,  with  an  elevation  of  5000  feet. 
The  1936  season  was  spent  at  Indian  Gardens,  22  miles  east  of  Payson,  in  an 
area  directly  beneath  the  rim  of  the  plateau  (here  called  the  Mogollon  or 
Tonto  Run).  No  specimens  were  found  in  this  well  forested  area,  though 


TREE  FROGS:  HYLIDAE  361 

frogs  were  found  on  the  plateau  above  on  several  trips.  Hyla  eximia  was  not 
listed  for  the  Grand  Canyon  National  Park.  .  .  . 

"Judging  from  the  small  number  of  specimens  found,  Williams  is  probably 
the  western  edge  of  this  frog's  range.  Although  the  summer  rains  were  few 
in  1937,  they  were  heavy  enough  to  have  brought  out  all  individuals,  but  very 
few  were  found.  Many  suitable  ponds  were  completely  barren.  Continuing 
east  numerous  specimens  were  found  along  the  rim.  At  Hart  Canyon,  which 
is  50  miles  south  of  Winslow,  and  at  Pinetop  and  McNary  the  numbers  in  the 
summer  ponds  were  very  great"  (W.  L.  Chapel,  Ariz.,  1959,  p.  225). 

Habitat:  "Before  and  after  the  breeding  season  the  frogs  are  found  occasion- 
ally throughout  the  forest.  Clever  camouflage  and  ventriloquism  aid  them  in 
escaping  detection.  They  are  found  on  the  ground  in  damp  places  and  in  the 
trees.  Twice  the  author  found  several  which  had  been  jarred  loose  when  a 
tree  was  felled.  In  one  of  these  instances,  a  frog  fell  from  the  top  of  a  tree 
about  75  feet  high.  On  sultry  days  they  were  heard  calling  from  the  trees" 
(same,  pp.  225-226). 

For  breeding  they  seek  large,  shallow,  grassy,  rain-water  pools,  permanent 
ponds,  new  ponds,  brooks,  even  wells. 

"It  is  a  species  apparently  adapted  to  semidescrt  conditions"  (E.  H.  Taylor, 
Ariz.,  1938,  p.  439). 

Size:  Adults,  1-1%  inches.  Males,  24-44  mm*  Females,  24-48  mm. 

General  appearance  and  Color:  At  first  gl.mce,  these  frogs  look  like  Ander- 
son's tree  frogs  (Hyla  andersonii),  green  Pacific  tree  frogs  (Hyla  regtlla),  or 
large  chorus  frogs  (Psettdacni).  They  are  bright  green  with  a  dark  purple 
or  black  line  from  the  snout  through  the  eye,  broadening  over  the  tympanum 
and  extending  halfway  along  the  sides.  This  vittal  stripe  may  be  broken  into 
groin  spots,  is  slightly  margined  above  with  white,  on  the  sides  is  irregular  in 
outline,  and  may  be  lighter  and  bronzy  in  the  center.  There  are  two  conspic- 
uous linear  black  spots  extending  backward  from  the  rump,  occasionally 
paired  dark  spots  on  the  forward  part  of  the  back,  but  no  marks  between  the 
eyes.  The  groin  is  conspicuously  greenish  orange  or  old  gold,  as  is  the  rc.ir 
of  the  femur,  which  is  unspotted.  There  arc  irregular  spots  or  bars  on  the 
tops  of  the  arms  and  legs.  There  are  irregular  dark  borders  on  the  upper  and 
lower  jaws  ending  in  a  dark  spot  at  the  shoulder.  The  throat  of  the  male  is 
dull  greenish  tan,  that  of  the  female  white. 

"This  species  is  related  more  closely  to  H.  regilla  and  H.  lajrentzi  than  to 
the  typical  H.  eximia.  From  the  former  it  differs  in  having  a  smooth  rather 
than  pustular  skin,  and  in  having  a  longer  leg,  the  tibiotarsal  joint  reaching 
the  tip  of  the  snout  or  beyond,  instead  of  to  the  region  of  the  eye.  The  webbing 
of  the  toes  is  somewhat  less  and  the  diameter  of  the  tympanum  is  greater  than 
half  the  diameter  of  the  eye;  the  toes  and  fingers  are  wider  with  somewhat 
wider  pads. 


362  HANDBOOK  OF  FROGS  AND  TOADS 

"From  H.  lajrentzi  it  differs  in  having  the  webbing  between  the  toes  some- 
what less  with  narrower  fingers  and  toes,  larger  choanae,  a  shorter,  blunter 
snout,  somewhat  deeper  in  front  ot  nostrils.  The  front  edge  of  the  tibia  is 
heavily-spotted  with  brown,  instead  of  having  it  blackish  with  a  cream-white 
or  silver  line  which  is  continued  to  foot"  (E.  H.  Taylor,  Ariz.,  1938,  p.  439). 

Structure:  Skin  smooth  above,  granular  below;  a  fold  of  skin  across  chest; 
a  fold  across  the  base  of  the  throat  in  the  female;  fingers  and  toes  with  well- 
developed  disks;  the  fourth  toe  very  long;  fingers  slightly  webbed  at  the  base; 
toes  %  to  l/2  webbed;  tympanum  half  the  diameter  of  the  eye;  tibia  half  the 
length  of  the  body;  a  tarsal  fold  present;  inner  metatarsal  tubercle  present, 
but  outer  metatarsal  tubercle  present,  inconspicuous,  or  absent. 

"Description  of  the  type.  A  medium-sized  member  of  the  Hyla  eximia 
group.  The  snout  is  rather  truncate  or  bluntly  conical,  with  the  canthi  more 
or  less  distinct  but  rounded;  the  line  between  eye  and  nostril  somewhat  con- 
cave, sloping  obliquely  from  canthus  to  edge  of  lip;  diameter  of  eye  somewhat 
greater  than  distance  of  eye  to  nostril,  and  equal  to  distance  of  nostril  to  mid- 
dle of  upper  labial  border;  nostrils  below  edge  of  canthus,  the  distance  be- 
tween them  about  equal  to  their  distance  from  eye;  the  area  about  nostril 
slightly  elevated,  and  a  slight,  shallow  groove  present  between  nostrils;  diame- 
ter of  the  tympanum  is  contained  in  the  diameter  of  the  eye  slightly  more  than 
1.5  times;  the  distance  between  the  tympanum  and  eye  about  .65  of  diameter 
of  tympanum. 

"Tongue  broadly  cordiform  or  subcircular  with  a  very  slight  median 
emargination  posteriorly;  free  posteriorly  for  two  fifths  of  its  length.  In  males 
the  openings  ot  the  single  vocal  sac  are  lateral  to  the  tongue  and  much  elon- 
gate; tongue  papillae  not  prominent;  the  raised  prominences  bearing  the 
vomerine  teeth  are  large,  placed  slightly  diagonally  and  closer  to  each  other 
than  to  choanae;  they  arise  near  anterior  level  of  the  choanae,  but  do  not  reach 
their  posterior  level.  The  openings  of  the  mucous  glands  form  a  continuous 
groove  anterior  to  choanae;  latter  proportionally  large. 

"A  vestige  of  a  web  between  first  three  fingers,  but  practically  obsolete  be- 
tween outer  fingers;  disks  on  the  fingers  moderate,  only  a  little  wider  than  the 
toes,  the  widest  one  on  outer  fingers  equal  in  width  to  a  little  more  than  half 
the  diameter  of  tympanum;  first  finger  reaching  to  a  point  halfway  between, 
the  distal  subarticular  tubercle  and  the  terminal  disk  of  the  second;  the  distal 
subarticular  tubercles  large,  that  on  outer  finger  very  slightly  bifid  on  right 
side  (probably  abnormally);  a  slight  dermal  fringe  on  the  lateral  edges  of 
fingers;  fourth  finger  longer  than  second. 

"Legs  elongate,  the  limb  laid  forward,  the  tibiotarsal  articulation  reaches  to 
the  tip  of  the  snout  or  beyond  slightly;  when  limbs  are  folded  at  right  angles 
to  body  the  heels  overlap  about  two  millimeters;  terminal  disks  on  toes  not 
wider  than  digits,  distinctly  smaller  than  finger  disks;  a  well-defined  tarsal 


TREE  FROGS:  HYLIDAE  363 

fold;  a  prominent,  salient,  inner  metatarsal  tubercle,  its  length  in  the  first 
finger  length  about  two  and  one-half  times;  outer  metatarsal  tubercle  distinct, 
flattened,  lying  behind  the  anterior  level  of  the  inner  tubercle;  inner  toes 
webbed  at  base,  the  depth  of  web  from  one  fourth  to  one  third  the  length  of 
the  outer  toes;  the  web  between  the  three  outer  toes  incised  to  a  point  one  third 
the  distance  between  the  two  proximal  subarticular  tubercles  of  the  fourth 
toe;  supernumerary  tubercles  on  palm  and  foot  more  or  less  distinct.  (In  males 
the  large  tubercle  at  the  inner  part  of  the  base  of  first  finger  is  covered  with  a 
corneous  callosity,  usually  very  light  brown  in  color.)  Anal  flap  rather  wide, 
not  especially  modified;  no  axillary  web;  skin  on  body  relatively  smooth,  un- 
der magnification  one  observes  minute  corrugations,  more  evident  above  eyes; 
a  strong  skin-fold  across  the  breast;  ventral  surface  granulate,  the  granules 
on  the  anterior  part  of  abdomen  largest,  less  distinct  on  throat  and  chin;  gran- 
ulations prominent  on  median  ventral,  and  to  some  extent,  posterior  part  of 
thighs;  a  rather  thick  but  relatively  indistinct  fold  above  tympanum"  (E.  H. 
Taylor,  Ariz.,  1938,  pp.  437-438). 

Voice:  The  call  is  a  low-pitched,  harsh,  metallic  clack,  with  no  trill,  and 
consists  of  two  to  ten,  twelve,  or  more  notes  given  in  succession.  These  may 
be  speeded  up  toward  the  end  of  the  call.  The  vocal  vesicle  is  very  large,  single, 
and  transparent,  with  less  yellow  or  green  in  it  than  is  found  in  most  tree 
toads. 

"Fresh  rains  bring  out  a  renewed  chorus.  A  chorus  usually  lasts  two  or 
three  nights  and  then  quickly  thins  out  to  a  few  disconsolate  males  for  two  to 
four  more  nights.  A  chorus  continues  all  night,  with  the  greatest  volume  be- 
fore midnight.  Probably  new  individuals  make  up  most  of  each  new  chorus" 
(W.  L.  Chapel,  Ariz.,  1939,  p.  226). 

Breeding:  July  9,  1942.  These  tadpoles  range  from  28-40  mm.  in  length 
(average  32  mm.)  with  body  lengths  of  11-17  mm-  (average  13.5  mm.)  and 
tail  lengths  of  17-23  mm.  (average  19  mm.). 

Tadpole.  Hart  Canyon,  CCC  Camp,  Winslow,  Ariz.,  July  16-24,  J933-  Hotly 
dark  olive  or  deep  olive  clotted  with  closely  set  mignonette  green,  light  yellow- 
ish olive  or  dull  citrine  dots;  the  snout  is  almost  the  color  of  these  dots  with 
no  olive.  The  deep  olive  of  the  back  extends  along  as  a  rim  margin  on  the 
upper  edge  of  the  musculature.  There  is  a  tendency  for  a  similar  rim  on  the 
lower  edge  of  musculature.  The  middle  portion  of  the  musculature  is  olive 
lake,  olivc-ochcr,  or  old  gold.  The  hind  legs  as  they  begin  to  develop  have 
much  the  same  color.  The  throat  also  is  this  color.  The  sides  of  the  belly  have 
a  flesh  color  or  light  vinaceous-cmnamon  cast.  The  middle  of  the  belly  is 
white.  The  tail  crest  is  finely  dotted  on  the  back  with  some  tendency  toward 
reticulation  of  the  dots.  The  lower  crest  to  the  naked  eye  may  look  to  be  almost 
transparent  but  is  actually  covered  all  over  with  very  fine  deep  olive  dots. 
Pupil  rim  of  eye  is  salmon  colored,  succeeded  by  black  iris  flecked  with  light 


364  HANDBOOK  OF  FROGS  AND  TOADS 

viridine  yellow  and  flesh  color.  Two  transformed  at  15  and  13.5  mm.  They 
were  green  at  transformation  and  had  a  suggestion  of  a  dark  indefinite  area 
or  band  ahead  of  the  eye. 

"The  breeding  season  is  from  June  to  August,  but  in  any  year  the  dates  vary. 
The  determining  factor  is  the  heavy  summer  rains  common  on  the  Colorado 
Plateau  in  July  and  August.  These  storms  delimit  the  breeding  season.  The 
seasons  noted  were:  1933  Pinetop  and  Hart  Canyon,  7000  feet,  July  2-Aug.  9, 
heavy  rains;  1935  Pinetop,  7000  feet,  July  7~July  28,  normal  rains;  1937  Wil- 
liams, 7000  feet,  July  2-July  14,  subnormal  rains.  Breeding  is  intermittent 
within  this  period"  (W.  L.  Chapel,  Ariz.,  1939,  p.  226). 

Two  ripe  females  37.5  and  42  mm.  were  taken  at  Santa  Fe,  New  Mexico, 
June  1 8,  1874,  by  H.  W.  Henshaw. 

"The  migration  of  frogs  toward  fresh  rain-water  pools  begins  as  soon  as 
the  summer  rains  start.  .  .  .  The  numbers  of  specimens  present  indicate  that 
large,  grassy,  shallow  ponds  are  quite  common  in  the  typical  open  parks  in 
the  ponderosa  pine  forests  in  central  Arizona.  The  grass  provides  resting 
places  for  the  frogs.  In  open  water  a  few  may  swim  about,  but  the  majority 
remain  along  the  edge,  resting  on  the  bottom.  .  .  . 

"The  duration  of  the  egg  and  tadpole  stages  is  unknown.  Some  eggs  were 
tagged  near  Pinetop,  but  the  author  was  transferred  the  next  day.  No  eggs 
have  been  found  elsewhere.  Some  tadpoles  were  coralled  several  times,  but 
subsequent  freshets  removed  the  broods.  Many  choruses  and  many  tadpoles 
have  been  found  in  rapidly  flowing  brooks  but  no  eggs  have  been  found. 
Probably  the  egg  masses  either  are  laid  in  quiet  back-washes  or  are  washed 
into  them  to  hatch.  The  frequent  freshets  cause  high  mortality,  and  after  the 
rainy  season  as  the  brooks  dry  up  many  tadpoles  are  isolated  in  pools  and  are 
dried  up  with  the  water. 

"Frequently  in  small  pools  the  water  is  nearly  black  with  the  swimming 
tadpoles.  The  tadpoles  gather  in  very  shallow,  warm  side-pools  full  of  de- 
caying vegetable  matter.  They  also  gather  in  great  numbers  around  fresh  cow 
manure  and  apparently  feed  on  the  dissolved  and  softened  material.  In  areas 
where  this  species  is  abundant,  in  later  summer,  when  the  tadpoles  transform, 
the  ground  around  the  ponds  is  covered  with  out-going  froglets.  These  vary 
in  length  from  10  to  13  mm.  Many  leave  the  water  still  carrying  stubby  tails" 
(W.  L.  Chapel,  Ariz.,  1959,  p.  226). 

Nov.  27,  1945.  Again  Bill  Chapel  arrives  in  the  "nick-o-time,"  to  tell  us  that 
those  eggs  he  tagged  near  Pinetop  were  in  small  loose  masses — possibly  the 
size  of  a  small  teacup — loosely  attached  to  grass  stems  just  below  the  surface 
of  the  water. 

Journal  notes:  On  June  10  or  12, 1933,  as  Mr.  W.  L.  Chapel  was  starting  for 
Arizona,  we  showed  him  illustrations  of  Hyla  wrightorum.  On  July  6,  he 
wrote  from  Winslow,  Ariz.,  somewhat  as  follows:  "At  present  I  am  at  Los 
Burros  CCC  Camp  near  McNary.  The  last  few  days  we  have  had  considerable 


TREE  FROGS:  HYLIDAE  365 

rain.  Toads  are  quite  numerous  about  camp.  Today  we  were  working  at  Los 
Burros,  which  is  about  six  miles  from  Winslow.  In  a  swampy  place,  I  heard 
quite  a  chorus.  I  found  endless  numbers  of  green  frogs.  No  other  species  could 
I  find.  I  am  sending  these,  eight  of  them.  .  .  ."  On  arrival,  they  proved  to  be 
four  males  and  four  females  of  Hyla  wrightomm. 

July  9, 1942.  Put  up  at  Lake-O- Woods  Lodge,  Ariz.  Went  down  to  swampy 
area  below  the  swimming  pool  dam.  Presently  in  water  2-4  or  6  inches  deep 
and  among  grass  found  tadpoles  of  Hyla  wrightomm.  The  tails  are  quite 
spotted  with  blackish  specks  and  in  general  they  are  rather  attractive  tadpoles; 
some  with  two  legs,  one  or  two  about  to  have  forelegs  break  through.  When 
did  they  breed? 

July  n,  Lake-O- Woods  Lodge.  Today  Anna  and  I  went  along  a  drainage 
ditch  west  of  the  lower  pond  and  down  to  a  swamplike  area  below  a  swim- 
ming pool.  Took  a  lot  more  tadpoles  ot  Hyla  wrightorum. 

Authorities9  corner: 

R.  Kellogg,  Gen.,  1932,  p.  168.  E.  H.  Taylor,  Kans.,  1938,  p.  439. 

F.  W.  King,  Ariz.,  1932,  p.  99.  W.  L  Chapel,  Ariz.,  1939,  pp.  225-227. 


FAMILY  LEPTODACTYLIDAE 

Genus  ELEUTHERODACTYLUS  Dumfril  and  Bibron 

Map  25 

Mexican  Cliff  Frog,  Robber  Frog,  Barking  Frog 

Eleutherodactylus  aitgitsti  (Duges).  Plate  LXXVII. 

Range:  Guanajuato.  Probably  not  in  the  United  States.  In  1933  we  gave  its 
range  as  Jalisco  to  southern  Arizona  on  the  basis  of  the  following  quotation: 

"A  frog  of  the  genus  Eleutherodactylus  has  been  received  from  Mr.  Sam 
Davidson,  of  Fort  Worth,  Texas.  This  specimen  collected  by  Mr.  Davidson, 
October  i,  1927,  in  Madcra  Canyon,  Santa  Rita  Mountains,  Arizona.  Miss 
Doris  Cochran,  of  the  United  States  National  Museum,  to  whom  this  speci- 
men was  sent  for  examination  and  comparison,  very  kindly  allowed  Dr.  Rem- 
ington Kellogg  to  examine  it,  and  it  was  referred  by  him,  on  account  of  the 
peculiar  dorsal  spotting,  to  Eleutherodactylus  augusti,  a  Mexican  species  liv- 
ing in  the  state  of  Guanajuato,  Mexico,  rather  than  to  Eleutherodactylus  la- 
trans,  known  only  from  the  state  of  Texas"  (J.  R.  Slevin,  Ariz.,  1931,  p.  140). 

In  1942  Mr.  Slevin  showed  us  this  specimen  and  we  concluded  that  super- 
ficially it  appeared  to  be  E.  latrans.  After  we  saw  Mulaik's  material  of  young 
with  light  crossbands  from  Kerrville,  Tex.,  we  more  than  ever  held  this  view. 
Koster's  (Carlsbad,  N.  Mex.)  and  our  (Boerne,  Tex.)  material  also  show  the 
light  transverse  band.  (Plate  LXXX.) 

Habitat:  (See  "Authorities'  corner.'*) 

Size:  Adults,  2%~3  inches.  64.5-75.0  mm.  (Kellogg). 

General  appearance:  "Mocquard  concluded  that  E.  augusti  was  identical 
with  Cope's  E.  latrans  from  central  Texas.  Direct  comparison  of  Mexican 
specimens  with  the  cotypes  of  E.  latrans  does  not  entirely  confirm  this  assump- 
tion. Though  there  are  no  constant  structural  features  that  will  distinguish 
specimens  from  these  two  areas,  it  was  observed  that  in  E.  latrans  the  fourth 
toe  is  relatively  longer,  the  color  pattern  consists  of  fairly  closely  aggregated 
large  black  blotches,  the  sides  and  hinder  half  of  the  abdomen  are  faintly  areo- 
late,  and  the  skin  on  the  upper  parts  of  old  adults  is  stiff,  coarse,  and  areolate. 
These  two  forms  are  unquestionably  rather  closely  related.  An  immature  in- 
dividual from  Jalisco  and  an  adult  individual  (with  a  body  length  of  75  mm.) 
collected  by  Ruthling,  which  unfortunately  is  without  any  definite  locality, 
were  used  in  these  direct  comparisons.  The  skin  on  the  upper  parts  of  the  im- 
mature individual  from  Jalisco  is  much  more  tubercular  and  warty  than  that 


kOBBER  FROGS:  LEPTODACTYL1DAE  367 

on  the  Texas  specimens.  Juvenile  characters,  such  as  vestigial  postcephalic  in- 
tratympanic  dermal  fold  and  vomcrine  teeth  in  minute  clusters,  are  not  un- 
usual, but  the  presence  of  an  abdominal  disk  seems  rather  remarkable  for  so 
young  an  individual"  (R.  Kellogg,  Gen.,  1932,  p.  101). 

"Hylodes  Augusti,  A.  Duges,  notes  manuscript.  Aspect  heavy.  The  head  is 
so  large  that  the  body  is  very  short;  the  eyes  large  and  prominent.  The  tym- 
panum is  conspicuously  smaller  than  the  eye.  The  tongue  is  a  little  longer 
than  broad  with  a  slight  notch  in  the  rear.  Vomerinc  teeth  in  2  oblique  groups 
in  the  rear  of  internal  nares.  The  body  is  finely  roughened  on  back,  smooth  on 
lower  surface.  Coloration:  The  upper  parts  are  a  very  clear  yellow,  inferior 
parts  white;  the  lower  side  of  the  throat  is  marbled  with  pale  brown,  the  sides 
and  the  arms  are  bordered  with  brown.  The  top  of  head  and  shoulders  are 
brown  black,  lined  with  pale  yellow.  One  sees  across  the  back  a  clearly  defined 
band  spotted  with  brown.  This  hylodes  is  considered  very  rare.  M.  Duges  dis- 
covered it  at  Guanajuato,  Mexico  .  .  ."  (Brocchi  T.,'  Gen.,  1879,  pp.  19-24). 

Journal  notes:  June  23, 1934.  In  the  afternoon  after  a  rain,  Anna  and  I  went 
to  Madera  Canyon,  Ariz.  Here,  Mrs.  Dusenberry  said  Mr.  Sam  Davidson  of 
Fort  Worth,  Tex.,  stayed  with  her  most  of  one  summer.  A  woman  came  from 
California  Academy  of  Science  to  collect.  (Miss  McLellan,  whom  he  later 
married.)  They  collected  the  frog  about  October  i.  It  is  the  specimen  now  in 
Calif.  Acad.  Sci. 

June  26,  Madera  Canyon.  Started  for  old  Baldy  about  7:30  A.M.  Arrived  at 
the  lookout  about  1 130.  L.  N.  Sprung  of  Sonoita,  Ari/ona,  is  the  fire  watch- 
man. He  was  here  in  1925  with  his  cousin,  Kenneth  Putnam,  also  of  Sonoita. 
Last  July,  just  below  the  watch  station,  Mr.  Sprung  one  afternoon  espied  a 
queer  frog  on  a  rock  face.  He  showed  it  to  two  boys  who  bothered  it  to  sec 
it  hop.  It  was  "more  round  than  any  other  shape,"  and  after  hopping  on  the 
rock  face,  it  went  into  a  crevice.  It  was  probably  the  robber  frog. 

In  1942  we  measured  the  California  Academy  of  Science  specimen  from 
Arizona.  The  measurements  (in  mm.)  of  this  female  arc:  length  85,  head 
(angle  to  mouth)  22,  width  of  head  57,  snout  15,  eye  14.5,  intcrorbital  space 
5.5,  upper  eyelid  8,  tympanum  6,  intcrtympanic  space  25.5,  internasal  space 
7,  fore  limb  46,  first  finger  15,  second  finger  15,  third  finger  2$,  fourth  finger 
21,  hind  limb  from  axil  106,  tibia  38.5,  foot  with  tarsus  50,  foot  without  tarsus 
32,  first  toe  9.5,  second  toe  15,  third  toe  21.5,  fourth  toe  32,  fifth  toe  21.5,  hind 
limb  from  vent  114.  Forearm  much  developed,  brachmm  diameter  6,  antc- 
brachium  9;  suggestion  of  a  disk  on  venter;  has  fold  of  skin  across  head  from 
tympanum  and  one  on  side  of  body  from  rear  of  tympanum  to  groin,  there 
bending  inward  to  the  vent.  It  is  E.  latrans. 

Authorities1  corner: 
P.  Brocchi,  Gen.,  1879,  p.  53. 
F.  Mocquard,  Gen.,  1899,  pp.  160-163. 
R.  Kellogg,  Gen.,  1932,  p.  101. 


368  HANDBOOK  OF  FROGS  AND  TOADS 

In  1938  Taylor  (Gen.,  1938,  pp.  391-394)  described  a  new  form,  E.  cactorum, 
and  contrasted  it  with  E.  latrans,  E.  augustt,  and  £.  laticeps.  He  had  a  key 
difference:  "Tympanum  %  to  %  diameter  of  eye;  dorsal  surface  smooth; 
Texas;  90  mm.  .  .  .  £.  latrans  Cope.  Tympanum  scarcely  more  than  one  half 
diameter  of  eye;  Guanajuata;  75  mm.  augustt  Duges."  In  addition  he  gave 
a  table  of  measurements  for  the  three  species. 

Texan  Cliff  Frog,  Barking  Frog,  Robber  Frog,  Rock  Frog 
Elctttherodactylus  latrans  (Cope).  Plates  LXXIX,  LXXX;  Map  25. 

Range:  Waco  to  Helotcs  to  Del  Rio,  Tex.,  into  Coahuila  (Schmidt  and 
Smith,  1944)  through  New  Mexico  to  Santa  Rita  Mountains,  Ariz.  We  had 
reports  of  it  at  Devil's  River,  Mertzon,  Hondo,  and  Frio,  Tex.  R.  D.  Quillin 
reported  it  from  Segovia,  Medina  Canyon,  Boerne,  Bracken,  San  Marcos, 
Concan,  and  Leakey,  Tex. 

"Eight  specimens  from  Sacaton,  five  miles  south  of  Cuatro  Cunegas.  All 
were  caught  in  mouse  traps.  Mr.  Marsh  states  that  search  for  specimens  by 
day  and  night  failed  to  discover  them.  They  apparently  got  into  his  traps  just 
before  dawn"  (K.  Schmidt  and  D.  W.  Owens,  Tex.,  1944,  p.  100). 

W.  Chapel  tells  us  (1945)  that  he  caught  a  good-sized  frog  with  a  promi- 
nent ventral  disk  which  he  thought  must  be  E.  latrans.  Alas,  he  lost  it.  He 
found  it  in  Parker  Canyon,  northeast  of  Roosevelt  Reservoir  in  central  Ari- 
zona. It  is  worthy  of  note  that  there  are  limestone  outcrops  in  that  region. 

Habitat:  Limestone  ledges  of  the  cliffs  that  front  the  Edwards  Plateau. 
They  have  also  been  reported  in  caves,  under  stones  and  other  cover,  in  rock 
walls  of  canyons,  rock  masses  in  mountains,  or  rocky  hillsides. 

Charles  T.  Vorhies  wrote  us  of  his  experience  in  Madcra  Canyon,  Arizona, 
"I  am  still  sure  of  one  thing  and  that  is  that  I  have  never  heard  such  notes 
from  any  other  species  of  frog  or  toad.  However,  the  specimen  I  captured  was 
not  on  a  cliff  but  merely  on  a  steep  rocky  hillside  and  was  sitting  under  a  j ul- 
ting  rock." 

Size:  Adults,  i%-3 %  inches;  48-90  mm. 

General  appearance:  This  is  a  large,  short-bodied,  squat  frog  with  extremely 
powerful  forearms  and  long  outer  fingers.  Its  fingers  and  toes  have  expanded 
tips  and  prominent  tubercles  on  the  under  surfaces.  The  loreal  region  is 
oblique.  The  skin  is  smooth.  The  back  may  be  light  drab  to  greenish. 

The  young  and  half-grown  may  be  greenish  on  the  back  and  have  a  light 
fawn,  unspotted,  transverse  band  across  the  middle  back. 

Color:  Male.  Helotes,  Tex.,  Feb.  21,  1925.  Background  of  back  light  drab 
shading  into  pale  drab-gray.  Light  area  down  middle  of  back  is  pale  drab- 
gray  or  pale  smoke  gray.  Spots  on  back  and  upper  parts  of  arms  and  legs  are 
natal  brown  or  army  brown;  sides  and  sides  of  belly  vinaceous-buflf;  legs  and 
arms,  hands,  and  toes  with  light  vinaceous-fawn  or  pale  vinaceous-fawn;  tym- 


ROBBER  FROGS:  LEPTODACTYLIDAE 


369 


Plate  LXXVII.  Eleutherodactylus  au- 
gusti.  1,3.  After  Brocchi  (X%)-  2.  After 
Brocchi  (XO-  4-  After  Mocquard 


Plate  LXXV11L  Eleutherodactylus  ri- 
cordii  planirostris.  1-4,6.  Adults  (X1)-  5>7- 
Embryo  developing  within  the  egg  (X5)* 
[No  free  tadpole.!  8.  Larva  freed  from 
gelatinous  envelope,  3  days  older  than 
fig.  7.  9.  Female  (Xi%)-  (5»7~9-  Paintings 
by  R.  F.  Decker t). 


370  HANDBOOK  OF  FROGS  AND  TOADS 

panum  vinaccous-slate  with  a  ventral  spot  from  center  up  of  pale  smoke  gray 
or  pale  drab-gray;  color  on  rear  of  thigh  unicolor  and  is  the  color  of  back- 
ground. Middle  of  throat  and  ventral  parts  are  pale  pinkish  buff.  Eyelid  is 
Varley's  gray;  iris  kght  yellowish  olive  with  a  bronzy  cast. 

In  G.  W.  Marnock's  collection,  Baylor  University,  no.  2022  has  a  prominent 
transverse  band  across  dorsum  from  arm  insertion  to  arm  insertion.  Back  of  it 
comes  a  broad  transverse  area  of  olive-buff  outlined  behind  by  two  dark  spots, 
one  either  side  of  the  meson.  This  prominent  light  band  has  a  median  light 
dorsal  stripe  extending  forward  on  meson  to  between  the  eyes.  USNM  no. 
13633  (G.  W.  Marnock,  Helotes,  Tex.),  which  measures  48  mm.,  has  an  indi- 
cation of  this  median  line  between  the  eyes,  and  possesses  a  rather  spotted  back. 

Half-grown  frog.  Cave  Without  a  Name,  Boerne,  Tex.,  from  V.  L.  Cock- 
rell,  Aug.  14,  1942.  General  color  of  snout  region,  area  across  shoulders,  and 
rear  of  back  is  deep  mouse  gray  heavily  spotted  with  large  spots  of  dark 
quaker  drab.  The  area  between  the  eyes  is  vinaceous-fawn  with  3  or  4  promi- 
nent black  spots.  Across  the  middle  back  ts  a  light  area  of  fawn  with  no  spots. 
The  tympanum  is  natal  brown.  The  femur  has  3  bands,  tibia  3,  metatarsus  2, 
antebrachium  i  of  army  brown,  the  interspaces  being  avellancous.  The  for- 
ward half  of  the  venter  is  pure  white  with  solid  pigments;  the  lower  belly, 
underside  of  femur  and  tibia,  and  inner  side  of  metatarsus  have  no  pigment 
and  the  flesh  showing  through  gives  appearance  of  light  vinaceous-lilac.  In 
the  eye  is  a  thin  pupil  rim  of  mars  orange,  the  base  a  deep  mouse  gray  almost 
obscured  by  dots  of  buff-yellow. 

Structure:  Male  with  forearm  best  developed,  brachium  little  developed; 
two  solar,  two  palmar  tubercles;  head  wide  and  flat;  muzzle  projects  beyond 
lower  jaw;  vomerinc  teeth  in  two  short  raised  patches  between  the  inner  nos- 
trils; tympanum  slightly  deeper  than  wide;  tongue  slightly  nicked  behind; 
fold  of  skin  from  eye  over  tympanum  almost  to  arm  insertion;  central  abdo- 
men surrounded  by  a  circular  fold  of  skin;  disks  transverse. 

The  enlarged  forearm  may  come  with  age.  Strecker's  no.  2022  (36  mm.) 
has  the  forearm  not  so  proportionally  larger  than  brachium  as  it  is  in  74-mm. 
and  77-mm.  specimens.  This  same  specimen  has  not  the  ventral  disk. 

In  some  alcoholic  specimens  there  seems  to  be  a  fold  of  skin  across  top  of 
head  between  the  tympani.  The  fold  bows  slightly  forward. 

Voice:  At  a  distance  the  call  was  certainly  a  bark,  but  as  we  climbed  the  hill 
and  came  near,  it  was  more  of  a  throaty  whttrr.  It  is  ventnloquial,  and  the 
location  of  the  calling  frog  back  under  the  ledge  doubtless  increases  the  vol- 
ume of  sound. 

"Roy  [R.  D.  Quillin]  returned  Sunday  [April  23,  1933]  from  a  trip  in  the 
Frio  Canyon  [Tex.],  and  the  barking  frogs  do  bark.  Mr.  Fisher  on  the  ranch 
said  that  the  frogs  made  so  much  barking  four  weeks  ago  that  he  was  awak- 
ened nights.  We  hear  them  every  evening  when  we  sit  on  the  front  porch  of 
the  ranch  house"  (letter  from  Mrs.  Ellen  Schulz  Quillm). 


372  HANDBOOK  OF  FROGS  AND  TOADS 

Mr,  Marnock  told  Mr.  Cope,  "During  the  winter  the  adults  are  very  noisy, 
the  rocks  resounding  in  the  evening  with  their  dog-like  bark.  The  noise  is 
supposed  by  the  country  people  to  be  made  by  lizards,  especially  the  Gcrrhono- 
tus  inferndis  which  occurs  in  the  same  region"  (E.  D.  Cope,  Tex.,  1878, 
p.  186). 

In  1925  this  last  notion  was  still  the  prevalent  one  in  Helotes,  for  we  were 
assured  that  it  was  the  "barking  lizard"  that  made  that  noise.  And  though  we 
caught  the  frog  that  was  "barking"  and  showed  it  to  the  doubters,  we  judge 
many  are  "of  the  same  opinion  still,"  for  since  our  return  we  have  received  a 
newspaper  clipping  with  a  picture  and  a  note  about  the  "barking  lizard." 

Feb.  20,  1925.  Heard  "barking"  near  the  store  at  Helotes.  Mr.  Fuller,  who 
keeps  the  store,  said  that  for  the  last  week  they  have  been  calling  but  very 
little  and  slightly  before  that.  Tonight,  we  went  to  Psettdacris  ornata  fork.  Air 
at  camp  7:45  P.M.  71°;  no  ornate  chorus  frogs  calling;  in  fact,  few  meadow 
frogs  calling.  Went  on  to  Helotes.  At  the  flat  crossing  right  near  Helotes  ham- 
let heard  a  "barking  lizard"  or  "frog."  Started  north  along  the  creek,  soon  dis- 
covered the  frog  was  way  up  on  the  hill  though  it  sounded  nearer  at  first. 
Went  across  open  field  back  of  the  store,  across  fence,  and  up  the  hill  among 
cedars,  rhus,  agerita,  prickly  pear,  and  yucca.  After  we  thought  we  were  near, 
we  discovered  it  farther  on.  It  is  not  startlingly  ventriloquial.  We  began  to 
suspect  it  was  going  to  be  in  a  shrubby  thicket.  When  we  neared  it,  it  did  not 
croak  5-15  times  in  succession  but  only  once  or  twice  at  a  time  with  longer  in- 
tervals between  croaking.  At  last  we  thought  we  had  found  the  ledge.  I  de- 
termined to  come  from  the  left  and  Anna  from  the  right  of  a  small  clump. 
I  halted  suddenly.  Four  or  five  feet  ahead  was  a  nice  diamondback  rattler 
asleep  or  coiled.  I  was  searching  frogs.  I  went  around  to  the  other  side.  Then 
a  long  interval.  Then  Anna  thought  she  heard  it  below.  Then  I  went  above 
where  I  thought  it  was.  Finally  it  croaked  between  us.  We  converged  only  to 
hear  it  in  another  direction.  At  last  we  determined  it  must  be  under  or  near  a 
rock  I  was  on.  Anna  below  tried  to  see  it,  and  I  from  above.  We  were  about 
to  give  up,  when  I  used  the  flashlight  on  it — and  saw  it  about  a  foot  under  a 
thick,  overhanging  rock.  Could  see  only  its  pulsating  throat.  Anna  put  flash- 
light on  one  side  to  prevent  its  escape  and  with  ruler  poked  it  out  far  enough 
for  me  to  get  a  hold  on  a  leg.  Then  the  game  was  ours.  It  truly  is  a  "robber* 
frog." 

A  card  from  A.  V.  Rutherford  of  Garner  State  Park  north  of  Uvalde,  Tex., 
written  March  22,  1947  states:  "A  man  here  has  caught  them  barking  high  up 
on  the  hills  under  ledges  of  rock.  He  heard  the  first  barking  this  year,  March 

15" 

Breeding:  Probably  they  breed  during  any  rainy  period  from  February  to 
May.  It  is  likely  that  the  large  eggs  are  laid  in  moist  or  rain-filled  cracks  or 
crevices  or  even  caves  in  the  rocky  cliffs  and  ledges  where  they  live. 

In  Sept.,  1930,  we  saw  a  large  female  (USNM  no.  10058)  which  was  col- 


ROBBER  FROGS:  LEPTODACTYLIDAE 


373 


lected  by  G.  W.  Marnock  at  Helotes,  Tex.  Dr.  Kellogg  had  discovered  that  it 
was  a  gravid  female.  It  measures  92  mm.  in  body  length.  The  egg  complement 
we  roughly  estimate  at  50  eggs.  These  eggs  have  the  largest  vitelli  of  any  frog 
eggs  of  the  United  States.  Measurements  of  four  vitelli  are  6.0,  7.0,  7.0,  and  7.5 
mm.  (/4-Kc  inch). 

It  seems  probable  to  us  that  development  is  in  the  egg  as  with  E.  ricordii, 
yet  it  is  possible  that  this  large  Elentherodactylns  may  have  a  method  diverse 
from  that  small  form.  We  were  in  Texas  in  a  very  dry  winter  and  spring,  but 
if  the  calling  frogs  bred,  we  could  not  find  their  tadpoles  in  the  creeks,  nor 
could  we  find  pools  of  water  near  the  ledges  that  might  harbor  them. 

The  likelihood  of  their  laying  their  eggs  in  a  stream  is  about  as  remote  as 
it  is  with  Camp's,  Marjiock's,  and  Ricord  s  frogs.  (See  P.  strecfyri,  "Authori- 
ties' corner,"  p.  280.) 

Journal  notes:  So  few  field  experiences  are  described  in  the  records  of  this 
frog  that  we  are  here  quoting  our  notes  at  length: 

Feb.  20, 1925.  Helotes,  Tex.  The  instant  I  had  it  in  hand  it  swelled  up  taut. 
When  I  reached  the  store  I  realized  I  was  sweating;  started  to  brush  sweat  out 
of  my  left  eye,  it  smarted  afterwards.  Some  cuts  on  hands  also  did.  It  affected 
mucus  of  mouth  slightly.  It  certainly  has  a  secretion.  It  can  bloat  out  bladder- 
like  more  than  any  other  form  I  have  ever  encountered,  as  much  so  or  more 
than  Hyla  gratiosa  or  some  Scaphioptts  species. 

Feb.  2i.  Heard  about  4:^0  P.M.,  air  temp.  71°,  one  robber  frog.  It  was  in  a 
crevice  of  limestone  on  Helotes  Hill.  Was  near  it,  too.  Just  back  of  the  first 
locality  of  last  night  is  a  Spanish  bayonet.  These  frogs  are  in  the  limestone 
ridges.  At  9  P.M.,  heard  none.  Air  70°.  Brec'/y  but  not  humid. 

Feb.  22.  In  photographing  male  1L  latrans,  it  would  wedge  itself  in  the  cell 
and  climb  right  out  of  the  jar.  When  1  had  it  in  hand,  it  bent  the  last  joint  of 
the  two  outer  digits.  These  have  expanded  tips.  With  these  bent  digits,  it  can 
pull  readily  and  strongly.  It  must  use  these  for  pulling  itself  along  or  to  pre- 
vent animals  from  pulling  it  out. 

March  18.  Tonight  about  9  P.M.,  temp.  70°,  heard  one  calling  on  Helotes 
Hill,  near  where  we  heard  one  on  Feb.  21.  Why  the  long  interval? 

March  20.  Tonight  about  9  P.M.  heard  one.  Temp,  about  65 °F. 

March  21.  Tonight  after  a  small  shower,  didn't  hear  a  robber  frog  on  the 
hills,  but  at  9:30  P.M.  our  captive  spoke  up  for  the  first  time  since  he  was  a 
captive.  Why? 

May  4.  Last  night  back  of  camp  we  heard  several  calling  until  midnight. 
Air  64°.  John  Barrara  (Mexican  ranch  hand)  said  they  called  vigorously  at 
5  P.M.  We  heard  several  north  of  Lee's,  also  east  of  Lee's.  We  heard  none  on 
Marnock's  hill  about  8  P.M.  We  rode  almost  to  Babcock  Road,  but  heard  few 
beyond  the  area  already  mentioned.  We  heard  one  on  hill  north  of  our  camp. 

.May  5.  On  hill  to  east  heard  robber  frog  a  few  times.  On  Marnock's  hill, 
heard  two  or  three,  then  they  ceased  calling.  Not  so  vigorous  tonight  as  last 


374  HANDBOOK  OF  FROGS  AND  TOADS 

night.  John  Barrara  says  they  get  down  to  the  creek.  The  one  on  our  hill  we 
thought  was  perhaps  moving  to  the  creek. 

May  9.  On  half  moon  of  rocks  and  hills  east  of  us  we  heard  many  robber 
frogs,  after  the  rain,  about  i  P.M.  Air  temp,  was  67°?.  This  morning  at  9, 1 
went  on  hill.  Worked  around  top,  saw  a  Holbroolya.  Then  came  back  at  top 
of  limestone  ledge,  and  then  back  at  base  of  limestone  ledge,  then  down  the 
canyon  in  the  half  moon.  I  did  not  see  a  water  pocket  of  any  permanence. 

July  4.  Storekeeper  at  Sheffield,  Tex.,  says  there  are  no  "barking  lizards" 
here.  About  85  miles  northeast  at  Mcrtzon,  they  occur.  Another  man  at  store 
said  they  are  also  near  Hondo.  This  accords  well  with  information  from  man 
at  Barton  Springs  Park.  We  were  told  of  their  being  at  Leakey,  Tex.,  in  the 
Frio  River  canyon. 

June  22, 1930,  San  Antonio,  Tex.  At  night  Roy  Quillin  and  I  went  to  John 
Classen's  ranch  to  listen  for  barking  frogs  where  he  and  Ellen  Schulz  Quillin 
hear  them  often.  Apparently  too  late  in  year  or  else  not  rainy  enough. 

June  23,  Helotes,  Tex.  Stopped  at  Mr.  Fuller's  store.  He  informs  me  he 
heard  many  on  the  hill  in  the  spring  but  not  lately. 

Feb.  26,  1942.  Went  to  California  Academy;  met  J.  R.  Slevin.  Glanced  at 
Eleuthcrodactylus  attgttsti.  It's  E.  latrans. 

Aug.  «;.  Went  south  of  Kerrville,  ideal  country  for  robber  frogs,  especially 
about  a  mile  or  less  north  of  Raven  Ranch  where  Stanley  Mulaik  got  a  few 
small  ones. 

These  young  arc  very  interesting  as  the  light  transverse  band  is  conspicu- 
ous, and  their  gray  color  in  preservative  is  suggestive  of  green  in  life. 

Later  in  day  went  to  Cave  Without  a  Name,  Mr.  Cockrell  guide.  His  wife 
said,  "Surely  there  are  salamanders  and  frogs  in  the  cave.  When  they  first 
opened  it  there  were  many  black  salamanders  on  the  walls  near  the  entrance, 
probably  our  cats  have  gotten  most  of  them."  When  Mr.  Cockrell  came  up, 
we  went  in.  He  said  he  saw  one  or  two  frogs  on  his  last  trip,  that  at  times  there 
were  three  or  tour  near  the  entrance.  In  the  morning  near  the  entrance  he 
found  one  of  them  in  a  water  pail. 

We  started  down  and  soon  in  a  round  pocket  in  the  wall  (pocket  6-7  inches 
deep)  was  a  beautiful  E.  latmns  headed  outward.  What  a  fine  sight  and  how 
nicely  niched  for  its  life!  Caught  it.  This  one  has  a  suggestion  of  the  light  band 
across  the  back. 

Authorities9  corner: 
J.  K.  Strccker,  Jr.,  Tex.,  I9o8c,  p.  59. 
J.  K.  Streckcr,  Jr.,  Tex.,  1922,  p.  15. 

"Enclosed  is  a  photo  of  a  frog  I  collected  about  12  miles  northwest  of  Carls- 
bad, N.M.  I  can't  find  any  description  in  our  library  that  fits  it  really  well.  For 
one  thing  the  coloration  is  quite  different.  It  comes  closest  to  the  Elcuthero- 
dactylns  latran*  collected  by  Marnock  which  you  describe  on  pp.  161-163  of 


ROBBER  FROGS:  LEPTODACTYLIDAE 


375 


Plate  LXXIX.  Eleutherodactylus  latrans. 
i.  Male  (X^)-  2.  Male  (X%).  3-  Male 


Plate  LXXX.  Eleutherodactylus  latrans. 
Upper,  i.  Young  adult,  from  W,  J.  Koster, 
near  Carlsbad,  N.M.  Center.  2,3.  Young, 
from  Stanley  Mulaik,  near  Kerrville,  Tex. 
r,  4.  Adult  from  Cave,  Boerne,  Tex. 


376  HANDBOOK  OF  FROGS  AND  TOADS 

your  Handbook.  The  following  is  a  description  that  I  made  after  the  speci- 
men had  been  in  formalin  for  two  days : 

"Most  of  the  dorsum  greenish,  darkest  on  head  and  neck,  venter  plain 
white.  Upper  jaw  whitish  with  vertical  green  bars.  A  double  whitish  line  ex- 
tends across  the  anterior  part  of  the  interorbital  region.  An  irregular  whitish 
line  extends  most  of  the  distance  between  the  interorbital  line  and  the  inter- 
scapular  region.  Body  with  numerous  dark  green  spots,  the  more  posterior 
ones  faintly  outlined  with  whitish.  Band  across  back  between  arm  insertions 
and  forelimbs,  ivory.  Forelimbs  with  a  bar  on  the  wrist,  forearm  and  a  faint 
one  on  the  upper  arm.  Hind  limbs  lighter  green  than  the  back,  with  darker 
green  bars,  these  more  or  less  outlined  with  lighter. 

"The  colors  were  not  greatly  changed  from  the  condition  in  life  except  for 
intensity.  It  was  strikingly  green  and  ivory  or  white  when  first  captured.  .  .  . 

"The  specimen  had  an  eventful  history.  It  was  taken  from  under  our  tent 
where  it  and  a  Cnemidophorus  apparently  sought  shelter  from  a  very  heavy 
and  violent  rainstorm  (3.48  inches  at  El  Paso  and  the  storm  was  still  con- 
tinuing when  we  broke  camp)  which  lasted  all  night  and  through  the  morn- 
ing. I  heard  short  barks  which  I  now  suspect  were  these  frogs  but  which  I 
then  thought  were  dogs.  I  brought  it  back  alive  and  when  I  recovered  I  tried 
to  identify  the  beast.  I  kept  it  in  a  deep  pan  of  water  to  keep  it  from  jumping 
whilst  I  was  looking  at  the  books  and  the  darn  thing  nearly  drowned.  As  a 
matter  of  fact  it  passed  out  and  I  gave  it  artificial  respiration.  It  died  during 
the  night.  After  it  was  preserved,  I  set  the  jar  on  my  work  table  next  to  the 
wall.  A  janitor  moved  the  table  for  the  first  time  in  three  years,  broke  the  jar 
and  threw  everything  into  the  basket.  When  I  returned  to  the  office  I  found 
the  specimen  in  the  basket  with  purple  "Ditto"  ink  spots  on  it.  It  has  since 
recovered.  You  see  why  I  did  not  send  the  specimen  right  off"  (letter  of  W.  J. 
Koster,  Albuquerque,  N.  Mex.,  Dec.  29,  1944).  See  condensed  account  by 
W.  J.  Koster,  N.  Mex.,  1946,  p.  173. 

"Though  it's  too  long  for  a  letter,  I'll  tell  you  my  story  of  what  I  have  for 
several  years  supposed  to  have  been  Eleutherodactylns.  In  1918,  while  engaged 
in  much  field  work  on  the  Santa  Rita  Experimental  Range,  I  had  my  family 
in  a  cabin  well  up  on  the  east  side  of  Madera  (White  House)  Canyon,  whence 
I  came  down  to  the  range  below  each  day.  This  was  the  last  week  in  June.  The* 
summer  rains  began  in  the  canyon  that  week,  with  an  afternoon  sprinkle, 
next  day  an  afternoon  light  shower,  and  next  day  with  a  rather  heavy  but  brief 
late-afternoon  rain.  About  dusk  a  loud,  raucous  calling  broke  out  in  the  can- 
yon. I  was  quite  new  to  this  country,  especially  to  summer  conditions,  and  a 
suggestion  that  it  was  'toads'  found  no  favor  with  me.  I  was  mystified.  I  went 
down  into  the  canyon,  only  to  find  the  sounds  came  from  the  opposite  side; 
climbing  up,  all  the  noise  hushed  up.  I  seated  myself  about  where  I  judged 
the  sounds  to  have  come  from.  After  several  fruitless  minutes  of  waiting,  I 
was  startled  by  a  sudden  'bark'  directly  behind  me.  Creeping  toward  an  over- 


ROBBER  FROGS:  LEPTODACTYLIDAE 


377 


hanging  rock,  there  was  just  enough  light  left  for  me  to  see  a  strange  'toad,' 
as  I  then  thought  it  was.  I  captured  it,  but  had  no  preservative,  and  I  lost  the 
specimen  by  escape  or  spoilage,  I  do  not  know  which.  Years  later,  when  I  first 
noted  a  report  of  Eleutherodactylus  in  Madera  Canyon,  I  guessed  I  had  seen 
and  heard  one.  I  still  believe  it  was,  and  have  hoped  to  run  across  it  again. 
But  I  have  never  since  been  in  Madera  under  similar  circumstances,  nor  have 
I  ever  heard  the  same  thing  under  other  conditions  when  up  there, 

"Last  summer,  having  some  field  work  out  that  way,  I  made  a  point  of  being 
up  there  on  a  late  June  day  when  both  clouds  and  calendar  indicated  a  pos- 
sible first  shower,  but  no  luck.  Again,  after  the  rains  had  begun,  I  explored 
pools  along  the  creek  in  the  same  section  of  canyon  without  result.  I'll  get  'em 
yet. 

"The  most  I  have  seen  of  the  Sierra  Ancha  has  been  in  late  fall — no  time 
to  find  these  frogs.  But  the  conditions  would  seem  to  be  as  good  as  in  the  Santa 
Ritas,  barring  its  being  further  north.  In  fact,  there  are  better-watered,  more 
precipitous  canyons  there  than  in  the  Santa  Ritas.  Do  you  think  I  had  Eleu- 
therodactylus*  (letter  of  C.  T.  Vorhies,  Jan.,  1946)  ? 

Ricord's  Frog,  Cricket  Toad,  Bahaman  Tree  Frog,  Pink-snouted  Frog 

Eleutherodactylus  rtcordii  planirostris  (Cope).  Plate  LXXVIII;  Map  25. 

Range:  Bahama  Islands,  Cuba,  and  Florida,  north  to  Duval  County.  In  the 
USNM  is  a  lot  of  two  specimens  (nos.  59419-20)  taken  at  Auburndale,  Fla., 
by  Wm.  Palmer.  In  recent  years  it  has  been  taken  as  far  north  as  Gainesville, 
Fla.,  by  O.  C.  Van  Hyning.  Each  of  these  records  reminds  one  of  Dr.  Bar- 
bour's  statement:  "It  has  been  taken  in  Florida  and  seems  to  be  spreading 
northward"  (Fla.,  p.  100).  In  1931  Dr.  H.  A.  Pilsbry,  while  collecting  pul- 
monate  gastropod  Liguus  on  or  about  Long  Pine  Key,  collected  several 
(F.  Harper,  Gen.,  1935).  In  1940  W.  G.  Lynn  and  in  1943  Lynn  and  J.  N. 
Dent  spoke  of  its  introduction  into  Jamaica,  where  it  is  now  widespread.  In 
1959  G.  A.  Skermer  of  Tampa,  Fla.,  published  some  of  his  experiences  with 
the  frog.  In  1944  C.  J.  Goin  had  specimens  from  Jacksonville. 

Habitat:  Leaf  piles  in  hammocks;  rock  piles;  fern  boxes;  under  walks,  logs, 
debris,  trash  piles;  limestone  heaps;  wells;  irrigation  banks.  Dr.  H.  A.  Pilsbry 
(1931)  found  them  under  loose  bark  in  the  hammocks. 

Size:  Adults,  %-iVr,  inches  (15-30  mm.). 

General  appearance:  It  is  small,  elongate,  similar  to  Syrrhophus  marnoc^ii 
but  possesses  two  long  transverse  curved  series  of  vomerine  teeth,  behind  the 
internal  nares.  These  are  delicate  little  frogs  with  the  head  as  broad  as  any 
part  of  the  body.  The  eyes  are  prominent,  beadlike,  bronzy,  and  black.  The 
snout  is  prominent,  truncate,  extending  slightly  beyond  the  lower  jaw.  The 
legs  are  long  and  slender.  The  fingers  and  toes  are  very  slender  with  terminal 
disks  and  prominent  palettes  at  the  joints  of  fingers  and  toes,  giving  a  saw- 


378  HANDBOOK  OF  FROGS  AND  TOADS 

tooth  appearance  in  latenil  aspect.  The  color  pattern  varies,  but  in  each  the 
coral  pink  snout  is  prominent. 

Color:  Adult.  Top  of  snout,  stripe  from  eye  to  hind  leg  insertion  along  dor- 
solateral  region,  interspaces  of  hind  legs  and  forelegs,  and  the  tubercles  along 
the  side  and  top  of  back  are  light  coral  red  to  dragon's-blood  red  or  terra  cotta. 
On  top  of  snout  is  a  dark  area  which  joins  with  the  dark  area  of  the  dorsum, 
which  normally  starts  about  on  level  with  a  line  drawn  from  middle  of  one 
eye  to  middle  of  other.  This  dorsal  color  is  brownish  olive,  snuff  brown.  In- 
distinct bars  of  the  same  color  on  femur,  tibia,  and  forearm  or  these  bars  and 
the  markings  on  the  toes  may  be  black.  The  upper  jaw  and  also  the  lower 
jaw  are  spotted  with  alternating  black  or  carob  brown  or  warm  blackish 
brown  spots.  A  line  of  the  same  color  over  tympanum  to  arm  insertion.  This 
line  sometimes  continuing  from  arm  insertion  along  side  as  more  or  less 
broken  dark  line.  Sometimes  from  axilla  to  groin  there  is  an  irregular  line  or 
row  of  dark  spots  parallel  with  the  upper  line  just  described.  Front  of  femur 
with  two  or  three  prominent  black  spots  and  several  such  spots  on  the  front 
of  femorotibial  articulation.  A  horizontal  line  of  same  color  from  arm  inser- 
tion halfway  down  the  brachium.  The  under  parts  are  white  and  heavily 
punctate  with  fine  brownish  olive  dots.  A  mid-ventral  blood  vessel  plainly 
shows  through  the  skin  of  the  belly.  The  alternating  areas  on  upper  and  lower 
jaws  are  white.  The  tympanum  may  be  almost  white  or  with  a  wash  of  coral 
red.  In  front  of,  below,  and  behind  eye,  below  tympanum,  and  along  sides 
are  spots  of  greenish  yellow.  Iris  is  principally  coral  red  to  dragon's-blood  red 
with  vermiculations  of  darker  color.  The  iris  rim  is  almost  scarlet  or  scarlet- 
red. 

Young.  The  conspicuous  thing  about  the  young  one  seems  to  be  the  light 
coral  red  triangular  snout  patch  ahead  of  the  eye,  the  same  color  on  fore  and 
hind  legs  and  a  patch  of  same  along  either  side.  In  most  of  the  young  there 
seems  to  be  an  appearance  of  triangular  spot  between  the  eyes,  the  base  of  the 
triangle  extends  between  the  eyes,  and  may  be  truncate,  marginate,  or  concave. 
The  rear  portion  of  the  triangle  oftentimes  and  generally  merges  with  the 
dorsal  band,  which  may  be  scalloped  along  its  sides.  It  is  a  beautiful  little 
creature  with  its  alternation  of  dark  and  light  spots  on  upper  and  lower  jaw 
and  with  its  big  bronzy  eye.  The  legs  appear  barred.  There  is  a  slight  ridge  ofc 
skin  down  the  middle  back — from  near  tip  of  snout  to  vent.  The  skin  of 
upper  surface  is  roughened  and  is  areolate  on  belly  and  hind  legs. 

Structure:  Subgular  vocal  sac;  tympanum  %  size  of  eye;  tongue  elongate 
oval,  slightly  nicked;  heel  reaches  orbit  or  even  snout;  lower  jaw  with  a  me- 
dian tubercle  fitting  into  a  median  notch  in  upper  jaw;  two  metacarpal  and 
two  metatarsal  tubercles. 

"Head  as  wide  or  wider  than  the  body,  longer  than  broad;  the  lateral  out- 
lines curved;  the  end  of  the  muzzle  abruptly  truncated.  Ostia  pharyngea  oval. 


ROBBER  FROGS:  LEPTODACTYLIDAE 


379 


Vomerine  teeth  in  two  long  curved  series,  which  commence  behind  and  oppo- 
site to  the  external  border  of  inner  nares,  they  are  separated  by  a  considerable 
space  medially.  Tongue  elongate  oval,  slightly  nicked.  A  subgular  vocal  sack. 
Tympanum  half  the  size  of  the  eye.  Skin  smooth  above  and  below;  sides  ru- 
gose. Heel  reaching  the  orbit.  Digital  palettes  small.  Two  metacarpal,  two 
metatarsal  tubercles.  Brachium  longer  than  or  equal  to  antebrachium.  .  .  . 
A  single  specimen  from  Key  West,  Florida,  is  in  the  National  Museum.  Its 
proper  habitat  is  Cuba"  (E.  D.  Cope,  Gen.,  1889,  p.  318). 

Voice:  "Its  twittering  call  can  be  heard  from  hammocks  as  well  as  dry  pine 
land,  after  showers  during  April,  May,  June,  July,  and  August"  (R.  F.  Deck- 
ert,  Fla.,  1921,  p.  22).  "Their  call,  which  they  gave  at  night  from  the  terrarium 
was  a  faint  'put  put' "  (E.  R.  Dunn,  Fla.,  1926,  p.  155).  "Their  chirping  notes 
are  a  common  sound  after  dark  and  on  cloudy  days"  (O.  C.  Van  Hyning,  Fla., 
1935,  p.  4).  "Some  five  years  ago  my  attention  was  attracted  by  a  trilling  sound 
which  seemed  to  come  from  a  cluster  of  ferns  around  a  palm.  At  first  I  thought 
it  was  some  new  cricket  or  grasshopper,  but  very  careful  search  and  watching 
did  not  solve  the  problem.  Although  I  have  lived  in  south  Florida  for  forty 
years,  I  had  not  heard  this  trill  before"  (G.  A.  Skermer,  Fla.,  1939,  p.  107). 

Breeding:  This  species  breeds  from  April  to  August.  Development  goes  on 
within  the  egg  to  adult  form,  there  being  no  tadpole  stage.  The  newly  hatched 
young  are  %-%  inch  (9.0-11.5  mm.)  in  size. 

"Ricord's  Frog  does  not  go  to  the  rain-pools  in  numbers,  as  do  the  other 
Salicntia.  Pairing  seems  to  take  place  on  land  during  rainy  weather,  in  dark 
places.  The  writer  has  so  far  failed  to  find  specimens  in  copula,  but  on  May  16 
two  batches  of  eggs,  containing  a  dozen  each,  were  found  in  a  depression 
filled  with  dead  leaves  and  leaf-mold  in  a  'hammock' "  (R.  F.  Deckert,  Fla., 
1921,  p.  23). 

"Ricordii  lays  19-25  eggs  in  vegetable  debris  in  woods,  .  .  .  yolk  being  about 
2  mm.  in  diameter,  and  the  outer  envelope  eventually  reaches  4  mm.  No  trace 
of  external  gills  could  be  made  out  in  either  species.  .  .  .  The  night  of  July  28 
a  batch  of  25  eggs  was  laid.  I  left  Soledad  on  July  30  and  returned  on  August  2 
and  found  that  a  second  batch  of  19  eggs  had  been  laid  in  my  absence. 

"The  eggs  of  the  28th  began  to  hatch  August  7,  making  a  period  of  eleven 
days.  By  August  n  all  were  out,  making  a  period  of  ten  days  from  the  2nd. 

"On  August  IT,  1925, 1  found  21  eggs  in  a  fallen  Bromeliad.  These  hatched 
six  days  after  (August  16)  and  as  soon  as  the  yolk  was  fully  absorbed  and  the 
adult  coloration  assumed  they  were  seen  to  be  ricordii'  (E.  R.  Dunn,  Fla., 
1926,  p. 155). 

"Presuming  that  this  frog,  like  others,  laid  its  eggs  in  a  lily  pond  near  its 
habitat,  I  did  not  give  thought  to  the  life  cycle.  While  looking  over  some  or- 
chids which  I  had  banked  with  sphagnum  I  found  a  cluster  of  what  appeared 
to  be  frog  or  toad  eggs.  I  had  not  seen  eggs  laid  in  this  way  before.  In  order 


380  HANDBOOK  OF  FROGS  AND  TOADS 

to  see  what  had  laid  them  I  lifted  the  cluster,  together  with  the  sphagnum  on 
which  they  were  deposited,  and  placed  them  in  a  glass  bowl  and  covered  the 
bowl  with  a  sheet  of  glass. 

"It  was  easy  to  see  that  the  eggs  were  normal,  for  the  embryo  could  be  seen 
moving  in  them.  In  about  seven  days  the  eggs  had  hatched,  the  jelly-like  cases 
lying  flattened  out  on  the  sphagnum.  After  very  close  inspection  of  the  sphag- 
num through  the  glass  I  saw  a  tiny  frog  not  as  large  as  a  house  fly  seated  on 
the  moss.  Its  appearance  was  exactly  that  of  the  mature  frog  I  had  previously 
found  so  I  knew  that  it  was  the  young  of  that  frog.  I  have  found  many  nests 
since  then,  all  made  on  the  ground  or  close  to  it  in  damp  shady  places"  (G.  A. 
Skermer,  Fla.,  1939,  pp.  107-108). 

"Mrs.  A.  N,  Dow  [of  Jacksonville,  Fla.]  found  eight  'small  white  eggs'  in 
a  flower  pot  in  her  yard  and  kept  them  until  they  hatched.  The  baby  frogs 
were  sent  to  me  for  identification,  and  proved  to  be  ricordii. 

"The  eight  individuals  vary  in  length  from  4.5  to  5.5  mm.  From  subadults 
from  Gainesville  they  differ  only  in  having  the  tympanum  slightly  less  dis- 
tinct, the  head  relatively  broader,  and  the  vomerine  teeth  less  well  developed. 
The  eight  specimens  are  now  catalogued  as  number  22557  in  the  collection  of 
the  Carnegie  Museum. 

"Mrs.  Dow  informs  me  that  the  locality  where  these  frogs  were  found  is  in 
a  residential  section  about  3  miles  from  the  business  center  of  the  city.  She 
found  the  eggs  in  a  flower  pot  which  had  been  left  undisturbed  in  a  flower 
bed  in  the  yard  for  several  months.  The  eggs  were  deposited  on  the  surface 
of  the  soil  and  when  Mrs.  Dow  first  discovered  them  she  placed  them  and  the 
accompanying  soil  in  a  jar  where  they  remained  until  they  hatched  about  a 
week  later.  She  sent  them  to  me  on  August  2, 1945,  when  the  frogs  were  'three 
or  four  days  old.'  In  answer  to  my  query  about  adults,  she  stated  that  she  had 
not  noticed  them  about,  but  that  a  neighbor  had  seen  frogs  of  this  description 
behind  the  houses  in  the  neighborhood"  (C.  J.  Coin,  Fla.,  1944,  P-  *92)' 

Apparent  newly  hatched  young  have  been  taken  in  the  months  of  June, 
July,  and  August.  The  two  following  records  are  of  young  apparently  not 
long  after  hatching:  one  specimen  (USNM  no.  36851)  taken  by  Prof.  C.  H. 
Eigenmann  on  July  3,  1905,  at  Santiago  de  las  Vegas,  Cuba,  measures  9  mm. 
and  others  (USNM  nos.  29823,  29825,  29838)  taken  by  Wm.  Palmer,  June  30,' 
1902,  at  Baracoa,  Cuba,  measure  11.5,  n,  and  11.5  mm.,  respectively.  In  the 
same  lot  were  some  specimens  15-20  mm.  in  length,  and  others  27-30  mm.  in 
length. 

We  have  seen  another  lot  (UMZ  64267)  from  Limestone  Ridge,  Soledad 
Cienfuegas,  Cuba,  7,  7.5,  9,  n,  and  19  mm.  in  length.  The  7  and  7.5  specimens 
must  be  near  hatching  size. 

Journal  notes:  (We  have  been  shown  the  actual  boards,  rocks,  and  vegetal 
cover  under  which  they  lived  and  the  actual  flower  pots  in  which  their  eggs 
have  been  taken,  yet  we  have  very  few  journal  notes  worthy  of  mention.) 


ROBBER  FROGS:  LEPTODACTYLIDAE  381 

Authorities9  corner:  "In  the  pine  country  [in  Dade  Co.,  Fla.]  the  writer 
has  found  it  under  heaps  of  limestone.  No  matter  how  dry  the  surrounding 
land  may  be,  in  the  center  of  these  rock-heaps  there  is  always  quite  some 
moisture,  and  all  kinds  of  creatures  find  hiding  places  there.  Many  specimens 
of  Ricord's  Frog  were  secured  in  rock  pits  under  quite  small  stones.  They  are 
difficult  to  capture  as  they  leap  with  lightning  rapidity  the  instant  they  are 
uncovered"  (R.  F.  Deckert,  Fla.,  1921,  pp.  22-23). 

"E.  ricordu  is  frequently  found  [in  Cuba]  along  with  cuneatus,  never,  how- 
ever, jumping  into  the  water.  Il  is  also  found  at  considerable  distances  from 
the  water. 

On  July  25, 1924, 1  caught  seven  from  a  colony  of  about  a  dozen,  which  was 
under  two  large  stones  and  the  fallen  leaves  of  a  royal  palm,  and  was  some 
fifty  yards  from  a  stream.  I  kept  five  of  these  alive  in  a  jar  with  some  vegetable 
debris"  (E.  R.  Dunn,  Fla.,  1926,  p.  155). 

"Eleiitherodactvlns  ncordn  is  a  Cuban  species  which,  it  has  been  assumed 
(Lynn,  1940),  has  been  introduced  accidentally  into  Jamaica  at  a  relatively 
recent  date.  It  was  first  recorded  from  Jamaica  by  Lynn  (1957)  who  took  it  at 
Montego  Bay.  Later,  E.  A.  Chapm  found  the  species  at  Hope  Gardens  in 
Kingston  and  it  was  supposed  (Lynn,  11)40)  that  this  indicated  two  separate 
points  of  introduction.  The  present  collections  however,  show  that  the  species 
is  wide-spread  m  the  island  and  it  may  be  that  it  has  been  in  Jamaica  for  a  long 
time.  We  took  specimens  at  Sandy  Bay,  Hanover,  at  Highgate,  St.  Mary;  near 
Port  Antonio,  Portland  at  Chapelton,  Clarendon  and  Mr.  L.  V.  Burns  has 
recently  taken  the  species  at  Hector's  River  on  the  border  between  Portland 
and  St.  Thomas"  (W.  G.  Lynn  and  J.  N.  Dent,  Gen.,  194$,  p.  259). 

"Since  Eleutherodactylits  ricordu  was  first  reported  in  southern  Florida  by 
Cope  (Bull.  U.S.  Nat.  Mus.,  (i),  [875:  31)  it  has  steadily  extended  its  range 
northward.  Carr  (Univ.  Fla.  Biol.  Sci.  Ser.,  Ill,  (i),  1940:  63)  gives  its  range 
as  continuous  in  south  Florida  northward  to  the  region  of  Melbourne  on  the 
east  coast  and  Clearwater  on  the  west  coast  with  a  separate  population  at 
Gainesville.  1  do  not  know  how  rapidly  the  southern  Florida  population  is 
spreading  northward,  but  the  Gainesville  population  is  certainly  expanding 
steadily  and  the  species  is  now  quite  common  several  miles  from  the  city  in 
localities  where  it  was  not  known  to  occur  a  few  years  ago"  (C.  J.  Coin,  Fla., 
1944,  p.  192). 

Shreve's  revision  (Fla.,  1945)  suggests  E.  planirortris  for  this  introduced 
frog. 

"I  therefore  restrict  the  name  ricordii  to  the  form  from  the  highlands  of 
Oriente,  Cuba,  which  has  a  spotted  dorsum.  The  color  is:  Above,  ground 
color  whitish  or  brownish  white,  heavily  spotted  with  dark  brown,  the  spots 
rounded  and  frequently  coalescing,  those  on  limbs  in  general  smaller.  In  some, 
the  spots  coalesce  to  such  an  extent  as  to  give  the  impression  of  marbling.  .  .  . 

"For  the  form  widespread  in  Cuba  and  the  Bahamas  hitherto  called  ricordii, 


382  HANDBOOK  OF  FROGS  AND  TOADS 

the  name  Eleutherodactylus  planirostris  (Cope)  is  available.  Its  type  locality 
is  New  Providence  Island,  Bahamas.  It  is  planirostris  therefore,  that  is  now 
present  in  Florida  as  an  introduction.  On  account  of  the  nature  of  the  differ- 
ences existing  between  ricordii  and  planirostris  (as  used  here),  it  seems  better 
to  use  the  two  names  in  subspecific  relationship,  although  in  the  material  at 
hand  there  is  no  real  evidence  of  intergradation  or  hybridization.  This  may 
be  shown  to  occur  eventually  as  there  is  some  evidence  of  hybridization  be- 
tween planirostris  and  the  species  casparit,  of  the  Trinidad  Mts.  in  Cuba, 
where  their  ranges  meet,  a  seemingly  parallel  situation"  (B.  Shreve,  Fla.,  1945, 
p.  117), 

Coleman  J.  Coin  (Fla.,  1947,  PP-  T~^)  has  just  issued  his  excellent  compre- 
hensive monograph,  "Studies  on  the  Life  History  of  Eleutherodactylus  ri- 
cordii planirostris  (Cope)  in  Florida."  It  emphasizes  particularly  habits, 
breeding,  development,  color  pattern,  and  distribution. 


Genus  LEPTODACTYLUS  Fitzinger 
Map  25 

White-lipped  Frog,  White-jawed  Frog,  White-jawed  Robber  Frog 

Leptodactyhts  lubialis  (Cope).  Plate  LXXXII;  Map  25. 

Range:  Mexico  to  extreme  southern  Texas  in  Starr  and  Hidalgo  counties. 
First  found  in  United  States  by  Dr.  E.  H.  Taylor  and  J.  S.  Wright  in  1931 
(Taylor,  Tex.,  1932).  In  1933  anc*  1934  it  was  held  to  be  Leptodactylus  albila- 
bris  (Gunther)  (Plate  LXXXI)  and  much  was  made  of  its  occurrence  in 
Puerto  Rico  and  on  the  mainland.  Dr.  Noble  in  1918  pointed  out  three  char- 
acters of  difference  between  the  mainland  and  insular  forms  but  frankly  ad- 
mitted that  no  criterion  is  constant.  K.  P.  Schmidt  in  1928  called  the  main- 
land and  insular  forms  very  close  but  was  not  able  to  state  that  they  are  iden- 
tical. 

Habitat:  Moist  meadows;  irrigated  fields;  drains;  gutters  in  towns;  beneath 
stones  and  logs.  Near  streams;  irrigation  ditches;  roadside  ditches;  in  bur- 
rows in  sandbanks  and  fields. 

"The  specimen  was  taken  late  at  night  hopping  about  on  the  table-like  top 
of  one  of  the  row  of  hills  just  north  of  the  Los  Olmos  bridge,  near  Rio  Grande 
City,  Tex.  The  earth  was  dry,  and  no  rain  had  fallen  for  a  period  of  forty-eight 
hours.  The  species  probably  shelters  itself  in  the  crevices  of  the  rocky  ledge 
which  surrounds  the  top  of  the  hill"  (E.  H.  Taylor,  Tex.,  1932,  p.  244). 

Size:  Adults,  i%-2  inches  (35-49  mm.). 

General  appearance:  This  is  a  small  dark-colored  frog  with  a  flattened 
pointed  head,  a  low  narrow  lateral  fold  and  a  cream-colored  labial  line. 
"Among  the  collections  made  in  Texas  during  the  summer  of  1931  by  J.  S. 
Wright  and  myself  is  a  small  leptodactilid  toad,  which  offers  something  of  a 
problepi  in  the  way  of  identification.  Through  the  kindness  of  Mrs.  Helen  T. 
Gaige,  Museum  of  Zoology,  Ann  Arbor,  the  specimen  was  examined  and  dis- 
sected, and  found  to  belong  to  the  genus  Leptodactyhts.  She  also  forwarded 
for  comparison  specimens  of  the  species  L.  albilabris  Gunther  from  Vera 
Cruz,  and  St.  Croix  Island,  which  seem  to  approach  this  form  in  many  char- 
acters. 

"From  these  two  forms,  which  differ  considerably  from  each  other,  I  note 
that  the  Texas  specimen  differs  in  the  absence  of  a  distinct  dorsolateral  fold, 
the  very  much  larger  choanae  (nearly  twice  the  area  of  those  in  the  compared 


384  HANDBOOK  OF  FROGS  AND  TOADS 

specimens),  the  presence  of  larger  triangular  bony  prominences,  on  which  the 
vomerine  teeth  occur,  and  their  separation  by  a  much  greater  distance  (more 
than  a  half  of  their  transverse  length) ;  the  very  narrow  lateral  fold  is  not  con- 
tinuous with  the  small  supratympanic  fold.  There  is  a  small  forward  exten- 
sion of  the  snout  to  form  a  'nose.'  The  dorsal  coloration  in  life  is  ash  gray  to 
clay  without  black  spots,  while  the  compared  forms  are  dark  blackish  brown 
with  well-defined  large  black  spots. 

"From  the  specimen  from  St.  Croix  (the  type  locality  of  Leptodactylus  albi- 
labris  Gunther)  I  note  that  the  tips  of  the  digits  are  distinctly  less  widened, 
in  fact  not  at  all  dilated;  that  the  head  is  longer,  more  pointed;  the  eye  rela- 
tively smaller;  and  that  the  ventral  brown  spotting  is  lacking. 

"From  the  Vera  Cruz  specimens  it  differs  in  having  a  flatter,  more  pointed 
head;  a  different  coloration.  It  also  differs  in  the  points  mentioned  above.  It 
agrees  with  both  forms  in  the  presence  of  a  well-defined  cream-colored  labial 
line,  and  the  presence  of  a  discoidal  disk  formed  by  a  skin  fold  on  the  abdo- 
men. 

"Cope  has  described  a  species,  L.  labialis,  which  has  small  choanae,  and  a 
paucity  of  dermal  folds,  and  L.  gracilis  with  a  projecting  muzzle  and  having 
dermal  folds. 

"Brocchi  has  described  L.  fragtlis  with  the  vomerine  teeth  in  arched  series, 
with  the  tympanum  nearly  twice  as  large  as  the  eye. 

"Remington  Kellogg,  who  is  reviewing  the  Amphibians  of  Mexico,  pro- 
poses to  place  labialis  Cope,  and  fragtlis  Brocchi  all  under  the  synonymy  of 
L.  albilabris  Gunther. 

"So  for  the  present  it  seems  that  this  Texas  specimen  must  rest  as  Lepto- 
dactylus albtlabris  Gunther  until  a  sufficient  series  of  specimens  arc  available 
to  determine  whether  the  separable  characters,  apparent  in  the  specimen,  are 
typical  or  due  to  individual  variation"  (E.  H.  Taylor,  Tex.,  1932,  pp.  243-244). 

Color:  "Above  rather  lead  gray  with  a  few  irregularly  scattered,  slightly 
dark,  grayish  blotches;  a  dark  vitta  from  snout  through  nostril  to  eye,  con- 
tinued behind  to  the  tympanum;  another  irregular  black  line  begins  above 
tympanum  and  follows  the  supratympanic  fold;  below  this  a  narrow,  well- 
defined  cream  line,  bordered  below  on  lip  by  an  irregular,  less  intense,  dark 
line  continued  to  angle  of  jaw;  a  few  black  spots  on  posterior  part  of  lower 
jaw;  arm  with  a  few  small  dark  spots,  and  a  few  also  in  axillary  region;  an- 
terior side  of  femur  with  a  dark  horizontal  line  and  small  spots;  upper  and 
posterior  surface  of  limb  spotted;  an  indistinct  cream  horizontal  line  borders 
the  granular  edge  of  the  under  part  of  the  femur,  below  which  are  a  few  small 
dark  spots;  under  surface  of  chin  through  abdomen  and  limbs  uniformly 
cream;  a  dark  broad  line  from  heel  across  foot.  The  measurements  are,  head 
and  body,  25  mm.;  foreleg,  13  mm.;  hindleg,  40  mm."  (E.  H.  Taylor,  Tex., 
1932,  p.  245). 

Structure:  Cope's  description  of  this  form  follows:  "Vomerine  teeth  in 


ROBBER  FROGS:  LEPTODACTYL1DAE 


385 


Plate  LXXXl.  Leptodactylus  dbilabris. 
i.  After  Brocchi  (X%)-  *•  After  Boulenger 
(X%).  3,4-  After  Schmidt  (X%)- 


Plate   LXXXII.   Leptoaaiiyim    w- — 
(X%)-  Adults,  from  Stanley  Mulaik,  Ed- 
inburg,  Tex. 


386  HANDBOOK  OF  FROGS  AND  TOADS 

transverse  series  behind  the  posterior  border  of  the  internal  nares;  toes  with- 
out dermal  border;  no  abdominal  discoidal  fold;  posterior  limbs  short;  end  of 
metatarsus  just  reaching  muzzle;  muzzle  short,  not  projecting;  teeth  behind 
choanae;  one  dermal  fold  on  each  side;  skin  rough;  below  white.  .  .  .  This 
small  species  belongs  to  that  division  of  the  genus,  in  which  the  toes  do  not 
possess  dermal  margin,  and  there  is  no  discoidal  fold  of  the  abdominal  integu- 
ment. Among  these  it  is  distinguished  by  the  shortness  of  series  of  vomerine 
teeth  and  the  paucity  of  dermal  plicae.  The  muzzle  is  acuminate  and  rather 
narrow,  but  not  projecting  as  in  C.  gracihs;  the  canthus  is  not  distinct.  The 
tongue  is  oval  and  a  little  notched  behind;  the  choanae  are  small.  The  diame- 
ter of  the  tympanic  disc  is  one  half  that  of  the  orbit.  The  heel  only  reaches 
the  orbit.  The  toes  are  not  very  long;  there  are  two  small  tarsal  tubercles  and 
a  narrow  tarsal  fold.  .  .  .  Color  chocolate  brown,  the  limbs  darker  cross- 
barred.  A  brilliant  white  band  extends  from  the  anterior  part  of  the  upper  lip, 
and  describing  a  curve  upwards,  bounds  the  orbit  below  and  descends  to  the 
canthus  oris,  from  which  point  it  continues  in  a  straight  line  to  the  humerus, 
and  ceases.  Inferior  surfaces  pure  white.  Length  of  head  and  body,  .020;  of 
head,  .007;  of  hind  limbs,  0.28;  of  hind  foot,  .013"  (E.  D.  Cope,  Gen.,  1877, 
p.  90). 

Taylor  states:  "The  snout  oval,  the  outline  broken  by  a  slight  pointed  pro- 
jection of  the  'nose';  the  head  measures,  length  11.5  mm.  to  angle  of  the  jaw; 
the  width  at  same  point  is  9.7  mm.;  canthus  rostralis  rounded,  the  ridge  con- 
tinued downward  in  front  ot  eye;  between  this  ridge  and  the  nostril  the  loreal 
region  is  distinctly  concave;  eye  moderate;  length  of  orbit  3.7  mm.,  the  dis- 
tance from  the  nostril  2.8  mm.  and  its  distance  from  the  tip  of  the  snout  4.2 
mm.,  tympanum  very  distinct,  rounded,  1.7  mm.  long  by  1.5  mm.  high,  sepa- 
rated from  eye  by  half  its  greatest  diameter;  a  slight  supratympanic  fold  from 
eye,  involving  upper  edge  of  the  tympanum  but  bending  down  some  distance 
behind  the  posterior  edge  of  tympanum,  labial  fold  distinct  posteriorly,  be- 
comes very  indistinct  anteriorly  and  terminates  below  nostril,  in  lateral  pro- 
file the  snout  extends  beyond  the  mouth ;  the  angle  of  mouth  reaches  directly 
below  tympanum;  the  distance  between  orbits  2.4  mm.;  of  upper  eyelid  width 
1.8  mm. 

"Skin  on  head  smooth,  showing  microscopic  corrugation;  the  dorsolateral 
fold  wanting,  represented  by  a  few  low  rounded  or  somewhat  elongate  pus- 
tules; a  sharply  defined,  very  narrow  lateral  fold  extends  from  above  arm  to 
groin;  above  this  a  few  elongate  or  rounded  pustules  low  and  indistinct;  below 
this  fold  the  skin  on  chin,  breast,  abdomen  and  limbs  smooth  except  for  a 
strongly  granular  distinctly  limited  area  on  ventral  femoral  region;  two  prom- 
inent granules  below  anus;  a  transverse  fold  across  the  breast  between  inser- 
tion of  arms  more  or  less  connected  with  a  lateral  abdominal  fold,  which 
curves  across  the  posterior  abdomen,  forming  a  disk;  a  small  diagonal  fold 
running  back  from  edge  of  disk  to  pubic  region;  limbs  not  especially  long, 


ROBBER  FROGS:  LEPTODACTYLIDAE  387 

the  tibia  12.5  mm.  is  distinctly  longer  than  the  femur  10.1  mm.;  from  heel  to 
tip  of  longest  toe  19  mm.;  an  elongate  sharply  defined  inner  metatarsal  tuber- 
cle, and  a  smaller  rounded  outer;  subarticular  tubercles  strongly  defined  save 
under  the  last  distal  joint;  heel  tubercular  with  a  distinct  fold;  rows  of  small 
tubercles  below  the  metatarsals;  tips  of  digits  not  dilated,  the  terminal  pha- 
lanx (bone)  pointed,  somewhat  clawlike;  no  webs  present  or  only  vestiges; 
two  large  palmar  tubercles;  an  elongate  inner  at  base  of  thumb,  and  a  larger 
rounded  basal  tubercle  with  an  anterior  extension;  first  row  of  subarticular 
tubercles  strongly  defined;  the  first  finger  longer  than  second  by  the  length 
of  the  last  phalanx.  .  .  .  The  measurements  are,  head  and  body,  25  mm.;  fore- 
leg, 13  mm.,  hindleg,  40  mm. 

"The  openings  of  the  choanae  are  well  back  from  tip  of  snout — 2.5  mm. — 
their  entire  outline  plainly  visible;  vomerine  teeth  on  two  triangular  raised 
patches,  separated  by  slightly  more  than  half  their  greatest  width;  they  are 
between  and  almost  wholly  behind  choanae  and  border  the  raised  eye  sockets1' 
(E.  H.  Taylor,  Tex.,  1932,  pp.  244-245). 

Voice:  "The  call  of  the  male  was  repeated  at  about  one  second  intervals 
at  the  height  of  rapidity,  and  resembled  the  plunk-plunk  of  a  drop  of  water 
falling  from  a  cave  roof  into  a  quiet  pool  below"  (S.  Mulaik,  Tex.,  1937,  p.  73). 

Breeding:  "The  finding  of  tadpoles,  transforming  young  and  a  frothy  egg 
mass  on  June  8,  1935,  immediately  following  heavy  rains  ten  miles  northwest 
of  Edinburg,  Texas,  closed  some  gaps  in  the  knowledge  of  the  life  history  of 
this  amphibian.  The  egg  mass  containing  86  yellow  eggs  was  found  in  a 
rounded  excavation  about  4  cm.  in  diameter  and  3  cm.  deep  at  the  base  of  a 
grass  hummock  a  foot  from  the  water  edge  and  several  inches  higher  than  the 
water  level. 

"These  eggs,  around  which  no  hyaline  sheath  could  be  discerned,  measured 
1.5  mm.  in  diameter.  Within  40  hours  after  the  eggs  were  laid,  the  young 
hatched.  And  were  practically  invisible  except  for  two  dark  eye  spots  and  the 
yellow  yolk  sack.  Several  hours  later  light  brown  pigmentation  appeared. 
The  emerging  tadpoles  measured  (in  mm.)  6.6  in  total  length,  2.3  in  body 
length  and  1.5  in  width.  Their  external  gills  remained  approximately  20  hours 
after  emergence.  They  were  apparently  two  branches  arising  near  the  same 
spot;  the  anterior  branch  was  the  larger,  measuring  about  i  mm.  in  length. 

"The  newly  hatched  young  rested  much  on  their  sides  or  even  on  their 
backs  on  the  bottom  of  the  aquaria  and  every  few  minutes  indulged  in  vio- 
lent squirmings  which  brought  them  toward  the  water  surface.  Twenty-four 
hours  after  hatching,  the  tadpoles  measured  8.1  mm.  in  total  length,  and  more 
pigmentation  was  noted.  On  the  second  day  the  young  were  observed  to  feed 
apparently  on  algae  and  diatoms,  and  their  measurements  were  as  follows 
(in  mm.) :  total  length  9.3;  body  length  3.3;  body  width  2. 

"On  the  third  day  measurements  of  u  mm.  in  total  length  were  recorded 
and  in  general  the  growth  appeared  rapid.  The  intestinal  coil  was  now  clearly 


388  HANDBOOK  OF  FROGS  AND  TOADS 

visible  through  the  transparent  abdominal  wall.  The  animal  food  provided 
was  not  accepted  before  the  eighth  day  at  which  time  the  tadpoles  began  to 
feed  upon  termites  offered.  Later  on,  as  they  developed,  the  tadpoles  readily 
fed  upon  other  insects  and  crushed  snails. 

"On  July  9  the  front  legs  of  one  of  the  tadpoles  emerged.  The  total  length 
of  this  specimen  was  32  mm.  or  6  to  8  mm.  less  than  the  same  measurement 
for  field  specimens  taken  at  this  stage.  The  tail  decreased  rapidly  from  this 
date  and  was  gone  by  July  n,  making  the  tadpole  stage  about  thirty  days. 
These  transforming  young  averaged  16.1  mm.  from  snout  to  vent;  rear  leg 
25.1  mm.;  hind  foot  12.6  mm.;  rear  leg  adflcxcd  brings  heel  to  rear  edge  of 
eye;  heel  to  tip  of  first  toe  6.6  mm.;  snout  to  angle  of  mouth  4.8  mm.;  tym- 
panum 2 1/2  times  into  eye. 

"Since  numerous  transforming  young  were  also  found  on  June  7,  and  since 
the  first  previous  heavy  rain  for  many  months  was  about  May  2,  the  tadpole 
period  for  the  field  specimens  was  judged  to  be  about  30  to  55  days,  about  the 
same  as  for  the  aquarium  raised  specimens.  These  young  were  hopping  about 
in  the  grass  at  the  edge  of  the  roadside  ditch  and  were  difficult  to  catch  because 
of  their  agility.  Though  adults  seemed  quite  common,  they  were  more  difficult 
to  take.  Males  when  calling  from  their  cavities  beneath  hummocks  of  grass, 
left  upon  discovery  and  quickly  sought  shelter  elsewhere.  It  is  believed  that 
the  males  constructed  these  cavities  to  be  used  by  the  females  for  the  deposi- 
tion of  the  eggs"  (S.  Mulaik,  Tex.,  1937,  pp.  72-73). 

Journal  notes:  May  i,  1934,  Edmburg,  Tex.  Mulaik  showed  me  two  frogs 
which  are  queer.  These  he  said  he  found  northwest  of  Edinburg.  They  were 
under  trash  and  mud.  By  digging  under  it  they  found  the  frog.  In  the  grass 
near  by  they  found  a  female.  They  sounded  like  sea  lions. 

May  }.  Trip  with  Stanley  and  Dorothv  Mulaik  northwest  of  Edinburg, 
Tex.  We  stopped  beside  the  road  near  a  ditch  or  pit  where  there  were  many 
grass  clumps.  Here  earlier,  when  there  was  some  water  in  the  pit,  they  had 
heard  the  "sea  lion"  call  of  I^eptodactylns.  In  one  clump  they  had  found  the 
male.  Today  we  dug  vigorously  but  it  was  too  dry  and  no  frogs  appeared. 

Authorities9  corner: 
K.  P.  Schmidt,  Gen.,  1928,  pp.  38-39. 
S.  Mulaik,  Tex.,  1937,  pp.  72-73. 


Genus  SYRRHOPHUS  Cope 


Map  25 

Camp's  Frog 

Syrrhophus  cum  pi  Stejncgcr.  Plate  LXXX1II;  Map  25. 

Range:  Brownsville,  Tex.  Lower  Rio  (irande  Valley. 

Habitat:  In  moist  earth  under  hoard  pile,  brick  pile,  stones,  or  similar 
shelter. 

Size:  Adults,  r's-i  inch  (15.0-25.5  mm.). 

The  largest  we  have  seen  is  a  ripe  female  25.5  mm.  A  collection  of  i;  adults 
taken  April  28,  1925,  in  Mr.  Camp's  board-pile  station  range  from  15-24  mm., 
the  average  19  mm.,  the  mode  19  mm.  A  series  of  9  taken  by  Mr.  ('amp 
(USNM  nos.  52^72-80)  range  from  i<V-2$  mm.  Another  series  of  ID  specimens 
taken  by  Mr.  Camp  (MCZ  nos.  10277-86)  range  from  15-25  mm. 

TABLE  OF  MKASURKMKNTS  (MM.) 


No. 

St\ 

Length 

1  lead  to  tympanum 

Head  to  angle  of  mouth 

Width  of  head 

Snout 

I-.yc 

Intcrorhital  space 

Upper  eyelid 

Tympanum 

Intertympanic  space 

Internasal  space 

Forchmb 

ist  finger 

2nd  finger 

$rd  finger 

4th  finger 

Hind  limb 


Syrrhophus 

Syirhophas 

cam  ft 

mainoit(ti 

USNM 

C.U. 

C.U. 

C.U. 

C.U. 

52291 

1277 

H7<> 

1271 

1382 

Ripe  9 

<J 

2O.O 

22.5 

25.5 

22.5 

26.0 

8.0 

8.0 

gjo 

<>-5 

10.5 

7-S 

6.5 

7.0 

8.0 

8.0 

7-5 

7-1? 

9-5 

8.0 

9.0 

4.0 

$-5 

4.0 

4-5 

5.0 

4.0 

4.0 

4.0 

4-5 

3-5 

3.0 

?•» 

i-5 

3-5 

I* 

2.0 

2.O 

2.0 

2-5 

2-5 

2.5 

2-5 

2-5 

2.0 

2.0 

7.0 

6.5 

7-5 

7-5 

7-5 

2.0 

2-5 

2-5 

2-5 

3.0 

13.0 

'?-5 

'3-5 

>5-5 

17.0 

M 

3.0 

3.0 

3.0 

*-5 

2-5 

3-5 

3-5 

3-S 

4.0 

4.0 

5.0 

5-5 

6.0 

6.0 

2-5 

4-5 

4-5 

4-5 

5.0 

27-5 

31.0 

31.0 

31.0 

34-o 

390  HANDBOOK  OF  FROGS  AND  TOADS 

Sytrhophus  Syt  rhophus 

campi  matnoclyi 

USNM        C.U.  C.U.  C.U.          C.U. 

Tibu                                         TO.O            10.0  11.5  12.0            12.5 

Foot  with  tarsus                        14.0            13.0  16.0  15.0            14.5 

Foot  without  tarsus                     9.5             9.0  10.0  8.5             7.5 

rst  toe                                          3.0             2.5  3.0  2.5             2.5 

2nd  toe                                         4.5             4.0  4.5  4.0             4.0 

?rd  toe                                         6  o             6.0  7.5  6.0             7.0 

4th  toe                                         9.5             9.0  10.0  8.5             7.5 

5th  toe                                         5.5             5.5  6.0  5.0             6.0  * 

*  I  hnd  foot  in  rather  poor  sh.ipc. 

General  appearance:  This  foim  is  very  similar  to  Syrrhophtis  marnoclyi, 
,uxl  belongs  to  a  Mexican  genus.  It  is  a  gravish  olive  frog,  with  scattered  dark 
spots  on  the  back,  with  a  dark  band  from  the  nostril  through  the  eye,  and 
with  dark  crossbars  on  the  legs.  The  skin  is  finely  granular.  The  nose  is 
pointed. 

Color:  Hrownsville,  lex.,  April  2S,  1^25.  Upper  parts  grayish  olive;  pale 
smoke  gray  from  eye  to  eye;  hair  brown  or  bcn/o  brown,  fuscous,  or  black 
vitta  in  tront  of  eye  meeting  fellow  on  snout.  This  line  goes  through  the  eye, 
over  the  tympanum,  above  the  shoulder  insertion  onto  body,  or  may  be 
broken  at  tympanum's  lear  edge  and  continue  backward  as  broken  spots. 
These  spots  may  go  almost  to  giom  or  stop  in  middle  of  side.  Sometimes  on 
kick  where  a  lateral  fold  normally  would  be  is  a  broken  longitudinal  black 
line.  On  the  back  aie  scattered  luscotis  black  or  hair  brown  spots.  Femur  with 
one  or  two  obscure  spots  on  front.  In  rear,  unitotm  light  grayish  olive;  tibia 
with  two  or  three  rathei  prominent  crossbars,  interspace  vctiver  green.  On 
sides  of  body  aie  elliptic  flecks  of  white  or  pallid  brownish  drab,  which  con- 
tinue to  below  e\e  or  tarthci  tot  ward;  one  or  t\\o  of  these  spots  are  just  belou 
and  behind  tympanum  and  sometimes  some  on  side  of  face  are  light  yellow- 
gieen.  Another  specimen  has,  starting  from  near  vent,  a  row  of  black  spots 
or  one  line  on  side  ot  back  hallway  to  shoulder.  Hind  legs  burred  and  spotted 
on  lemur  and  tibia  \\ith  black  and  Isabella  color,  which  become  white  or  pal- 
lid brownish  drab  spots  on  foot  and  toes.  Under  parts  of  breast  back  of  arm 
insertion  and  torward  part  of  bell\  arc  solid  pale  yellow-green.  Blood  vessel 
shows  thiotigh  its  veij  middle.  Lower  belly  and  rest  of  under  parts  are  pale 
piuphsh  vinaceous  to  light  purplish  vinaceous.  Forearm  is  spotted  with  white 
spots,  brachiurn  with  prominent  light  greenish  yellow  and  dark  spots  or  the 
light  green-\ellow  may  become,  in  some,  daik  olive-buff  or  deep  olive-buff. 
A  female  has  fe\\  if  am  dark  spots  on  the  back.  Ins  is  light  greenish  yellow 
above  and  deep  chrome,  with  one  vermiculated  line  of  brownish  through  it; 
in  the  rear  it  is  solid  black  below  and  in  front  it  is  black  with  light  greenish 
yellow  specks;  pupil  elliptical. 


ROBBER  FROGS:  LEPTODACTYLIDAE 


39* 


Plate    LXXXIII.     Synhophus     campi 
(X'%).  1-8-  Adults- 


Plate  LXXXIV.  Syrrtiopnus  marnw\u 
(Xi).  1-6-  Adults. 


392  HANDBOOK  OF  FROGS  AND  TOADS 

Structure:  Small,  delicate,  flat  in  body;  a  blood  vessel  visible  down  the 
middle  of  the  belly;  fingeis  and  toes  long  and  slender  with  prominent  tuber- 
cles, which  viewed  from  the  side  appear  saw-toothed;  forearms  well  devel- 
oped; tips  of  fingers  expanded  and  truncate;  toes  less  so;  at  least  one  light- 
colored  tubercle  just  back  of  angle  of  mouth,  and  near  the  lower  rear  margin 
of  the  tympanum;  often  a  collection  of  tubercles  in  this  region  or  an  oblique 
row  to  the  arm  insertion.  Wrist  extends  to  tip  of  muzzle.  In  alcohol,  the  sides 
and  upper  parts  of  hind  and  fore  limbs  seem  to  be  covered  with  little  round 
white  spots  amongst  the  punctae. 

Voice:  This  frog  gives  a  brief  cricketlike  chirp,  but  with  a  whistle.  It  is  not 
a  continued  call,  but  often  consists  of  only  one  or  two  notes  which  can  be  heard 
a  few  yards  away. 

April  28,  1925.  Early  in  the  morning  went  to  the  board  pile  station.  Heard 
one  chirp  over  in  the  west  shed,  one  or  two  more  chirping  in  a  crack  in  the 
brick  wall  near  the  ground.  Heard  several  in  the  board  pile. 

June  17, 1950,  Brownsville,  Tex.  They  are  a  dooryard,  front-porch,  or  garden 
friend,  cheerful  as  a  cricket.  The  note  may  be  one  ttcl{  or  two  or  three  ticl(s, 
these  well  measured.  Or  the  two  mav  be  rapid  and  close  together.  Sometimes 
after  these  comes  a  cricketlike  call. 

Breeding:  This  species  breeds  from  April  to  May.  The  eggs  arc  few,  6-12, 
and  very  large,  the  egg  yolks  about  Vis  inch  (3.0-3.5  mm.).  Larval  develop- 
ment is  probably  within  the  egg.  The  smallest  frogs  we  have  seen  in  collections 
range  %-%o  inch  (5.0-8.5  mm.). 

April  28,  1925,  at  Brownsville,  Tex.  We  secured  a  good  series  of  these  little 
frogs.  One  female  contained  very  large  eggs. 

May  5,  Helotes,  Tex.  This  evening  about  5:30  P.M.  I  looked  at  the  Browns- 
ville Syrrhophus  cumpi  to  compare  with  our  lone  5.  mtirnocfyi.  Lo  and  behold, 
a  male  was  holding  a  female  in  inguinal  amplexation.  The  fists  met.  One  fin- 
ger reached  across  the  thigh  and  the  other  across  the  lower  belly.  The  male 
is  more  spotted;  the  female  almost  without  spots. 

May  6.  Tonight  we  had  company.  When  I  showed  them  the  S.  campi  speci- 
mens, there  were  three  in  one  embrace,  one  above  another.  Each  of  the  upper 
two  had  an  inguinal  embrace. 

A  ripe  female  taken  April  28,  1925,  has  six  or  seven  very  large  ovarian  eggs 
well  advanced.  These  egg  yolks  range  from  5.0-3.5  mm.  in  diameter,  mosfof 
them  about  3  mm. 

The  very  few  large  yolked  eggs  and  the  very  small  frogs  seem  to  confirm 
the  opinion  of  Drs.  Stcjneger  and  Barbour  and  Messrs.  R.  D.  Camp  and  A.  C. 
Weed  that  the  larval  development  is  carried  through  in  the  egg. 

A  lot  of  10  small  frogs  sent  to  Dr.  Stejnegcr  by  Mr.  Camp  measured  6.5, 7.5, 
7.5, 9,  9,5,  9.5,  TO,  10,  11.5  mm.  Others  equally  small  were  seen  in  Mr.  Camp's 
collection.  The  American  Museum  has  3  specimens  (7.5-9.0  mm.).  The  Mu- 
seum of  Comparative  Zoology  has  18  small  specimens:  one  5  mm.,  one  6  mm., 


ROBBER  FROGS:  LEPTODACTYLIDAE 


393 


six  6.5  mm.,  seven  7  mm.,  one  7.5  mm.,  one  8  mm.,  and  one  8.5  mm.  (5-8.5  mm.) . 

Journal  notes:  April  27,  1925.  Mr.  Camp  of  Brownsville,  Tex.,  took  me  to 
the  type  locality.  It  was  in  a  small  back  yard  of  a  private  residence  in  the  center 
of  the  city.  In  the  rear  ot  the  house  was  a  laundry,  the  floor  of  which  was  close 
to  the  ground.  Against  the  rear  of  the  house  were  two  small  piles  of  brick, 
each  of  which  was  no  more  than  2-3  feet  across  and  i%-2  ft.  high.  The  ground 
was  moderately  moist  there.  Mr.  Camp  said,  "Move  these  bricks  and  you  will 
find  them."  He  warned  me  to  grab  quickly  but  I  was  not  quick  enough  for 
the  first  one.  I  had  turned  over  two  or  three  small  piles  of  bricks  and  become 
quite  discouraged  when  I  espied  another  under  the  shade  of  a  little  fig  tree. 
Presently  I  caught  my  first  one.  The  associates  of  these  creatures  are  sow  bugs, 
ants,  snails,  and  spiders.  Something  jumped  in  a  hole;  all  I  got  was  the  im- 
pression of  the  jump.  In  another  case  there  was  a  jump  under  a  board  which 
was  under  the  brick  pile.  Saw  12  or  15,  but  all  I  captured  were  four  adults — 
17  mm.,  20  mm.,  24  mm.,  24  mm.— and  a  half-grown  one  12  mm.  in  length. 
Only  a  keen  collector  would  have  discovered  this  new  species. 

April  28.  It  certainly  seems  as  if  they  come  out  to  the  edge  of  the  board  pile 
at  dusk  or  in  the  evening.  Under  one  square  board  i  ft.  or  i%  ft.  found  four 
at  the  edge  of  the  pile.  Do  they  feed  at  night ?  Someone  said  they  did  not 
want  to  lose  them  all  because  they  fed  when  mosquitoes  do  and  fed  on  mos- 
quitoes. 

April  29.  Went  to  the  board  pile  at  9:30  A.M. — mighty  hot.  Once  or  twice 
while  we  were  moving  the  pile,  we  heard  the  chirp  of  this  species.  Beneath  this 
pile  is  an  old  cistern,  now  partially  filled  with  bricks  and  dirt.  It  is  very  moist  in 
this  cistern,  but  no  water.  We  saw  two  or  three  frogs  in  amongst  the  bricks. 
We  moved  about  two-thuds  of  the  pile,  secured  eight  or  nine  frogs.  We  saw 
no  signs  of  eggs.  One  we  caught  rather  harshly  gave  up  a  lot  of  cockroach 
eggs.  These  he  wouldn't  need  to  feed  on  at  night.  We  didn't  find  any  of  the 
very  little  ones,  cither  in  the  board  pile  or  the  yard.  These  frogs  were  very 
quick,  leaping  the  instant  they  are  exposed.  They  arc  very  flat  and,  like  the 
robber  frog,  have  the  forearm  well  developed.  When  under  board  or  cover, 
they  often  crawl  along. 

June  16,  1930.  Walked  around  Brownsville  about  11-12  P.M.  Was  sure  the 
cheerful  chirp  I  heard  in  many  yards  was  Syrrhophns  campi. 

June  17.  About  5  P.M.  went  over  to  the  board  pile  station.  The  board  pile 
which  Camp,  Weed,  and  we  have  successively  moved  and  put  back  again 
is  gone,  but  Mr.  O.  P.  Hacker  who  owns  it  (corner  Washington  and  T5th 
Street,  Brownsville,  Tex.)  said  there  are  frogs  in  the  same  old  place.  Mr.  A. 
Holm  tells  me  this  area  was  General  Zachary  Taylor's  old  stables.  Went  into 
a  shed  near  the  old  place.  Began  turning  over  debris,  sticks,  wet  papers,  iron 
pieces,  and  clothes.  Under  these  the  little  frogs  were  revealed.  The  first  two 
I  lost.  Some  appeared  quite  "yellowish  green"  or  old  gold  with  spots.  They 
can  crawl  rapidly.  Most  of  the  specimens  were  quite  dark.  In  various  places 


394  HANDBOOK  OF  FROGS  AND  TOADS 

while  at  the  task  I  heard  their  pleasant  notes.  They  are  quick  to  find  a  crack 
in  the  wall  or  other  cover.  Where  the  old  board  pile  was,  now  bricks  are  in  a 
pile.  These  I  didn't  move  but  in  the  pile  were  frogs  chirping.  Took  several 
small  frogs.  One  very  tiny. 

June  19.  Mr.  Rebb's  palm  grove.  Arrived  at  7.  Palm  grove  flooded.  In  palm 
grove  are  plenty  of  S.  campi  near  its  unflooded  edge.  Others  sound  as  if  in 
deeply  flooded  parts.  Are  they  temporarily  on  boles  of  trees,  on  large  palm  fans, 
or  in  a  litter  of  palm  leaves  at  the  surface  ?  Surely  this  species  is  not  scarce.  It 
is  common. 

May  2,  1934,  Harlingen,  Tex.  Mr.  and  Mrs.  L.  Irby  Davis  say  they  have 
plenty  of  small  frogs  under  their  plant  boxes.  They  are  S.  campi.  Saw  several 
of  them  at  the  edge  of  their  rear  lawn. 

Authorities9  corner: 
L.  Stejneger,  Tex.,  1915,  p.  132. 

Gaige's  Frog 

Syrrhophits  gaigcae.  Map  25. 

Range:  "Type  from  the  Basin,  Chisos  Mountains,  lirewster  County,  Texas 
No.  27361  Field  Museum  of  Natural  History.  Collected  July  24,  1937,  by 
Walter  L.  Necker.  .  .  . 

"The  sixteen  paratypes  agree  well  with  the  additional  specimens  which  were 
taken  in  the  Basin  and  one  from  Pulham  Canyon  by  the  junior  author.  Ten 
additional  specimens  from  Juniper  Canyon  were  collected  by  Mrs.  Helen  T. 
Gaige  .  .  ."  (Schmidt  and  Smith,  Tex.,  1944,  P-  80). 

Habitat:  After  our  examination  of  the  Gaige  material  of  the  University  of 
Michigan  some  15  years  ago  we  wrote:  "The  Gaiges  |  Dr.  F.  K.  and  Mrs.  Helen 
Thompson]  became  very  familiar  with  this  species  while  they  were  collecting 
in  the  Chisos  Mountains,  Texas.  They  came  to  know  its  nocturnal  habits  and 
its  notes  and  collected  a  good  series  of  adults."  How  appropriate  that  in  1944 
the  authors  Schmidt  and  Smith  "named  |  this  frog  |  for  Mrs.  Helen  T.  Gaigc, 
in  order  to  associate  her  name  with  the  hcrpetological  exploration  of  the  Chisos 
Mountains,  in  which  she  had  a  pioneer  part." 

Size:  Adults  21-28  mm.  Mrs.  Gaige's  ten,  collected  15  years  earlier,  we  made 
21-31  mm.,  i.e.,  slightly  lower  than  our  and  Schmidt  and  Smith's  range  for 
S.  niarnoctyi. 

General  appearance:  "Closely  allied  to  Syrrhopus  marnocfyi,  from  which  it 
is  distinguished  by  smaller  size,  wider  head,  and  vermiculate  instead  of  spotted 
dorsal  pattern.  .  .  .  These  range  in  length  from  21  to  28  mm.,  compared  to  25 
to  35  mm.  in  marnoclyi;  the  ratio  of  width  of  head  to  body  length  is  0.40  in 
seventeen  gaigcae  and  0.36  in  twenty  marnocfyt.  The  paratypes  agree  with  the 
type  in  vermiculation  and  diflfusencss  of  the  dorsal  pattern"  (K.  P.  Schmidt 
and  T.  F.  Smith,  Tex.,  1944,  p.  80). 


ROBBER  FROGS:  LEPTODACTYLIDAE 


395 


Color:  "Pale  brown  above,  with  dark  brown  reticulation;  under  surfaces 
uniform  very  pale  yellow;  limb  with  broad  dark  crossbars"  (same). 

Structure:  "Syrrhopus  gaigeae  sp.  nov.  .  .  .  Description  of  type. — Habitus 
of  Syrrhopus  imtrnoclyt,  head  wider  than  body,  its  width  equal  to  the  distance 
from  snout  to  posterior  border  of  tympanum;  snout  long,  the  nostril  close  to 
the  tip;  canthus  rounded;  length  of  eye  much  less  than  its  distance  from  the 
nostril,  about  equal  to  the  diameter  of  the  tympanum,  which  is  very  distinct; 
heels  overlapping;  heel  reaching  midway  between  eye  and  tip  of  snout;  disks 
of  two  outer  fingers  much  wider  than  those  of  inner  fingers;  disks  of  toes 
small;  first  finger  about  equal  to  second;  metacarpal,  metatarsal,  and  subdigital 
tubercles  distinct;  skin  of  back  slightly  tuberculated;  a  rounded  gland  above 
the  insertion  of  the  arm;  no  glandular  fold  over  tympanum"  (same) . 

Journal  notes:  We  have  no  actual  specimens.  Once  in  the  Davis  Mountains 
we  were  sure  we  saw  a  Syrrhophus  disappear  under  a  huge  stone  and  in  the 
same  way  we  were  unsuccessful  in  Devil's  River.  Is  there  a  wide  gap  of  250 
miles  in  Syrrhophus  range  or  a  wide  gap  in  our  knowledge?  Again,  Edwards 
Plateau  to  Trans-Pecos  needs  attention. 

Authorities'  corner:  In  the  account  of  this  species  we  have  used  Schmidt  and 
Smith's  account  almost  exclusively  for  there  is  little  else  of  record.  They  write 
(p.  80): 

"The  new  form  is  admittedly  distinguished  from  marnocfyt  by  only  trivial 
characters;  it  is  believed,  however,  that  the  differences  noted  characterize  a 
well  isolated  population,  separated  by  some  250  miles  from  the  range  of 
mar noc kit,  and  that  these  differences  indicate  that  a  distinct  form  is  in  the 
process  of  evolution." 

About  1930  we  measured  two  of  the  University  of  Michigan  specimens,  each 
28  mm.  in  length,  and  compared  them  with  two  S.  marnocl{n  of  28  mm.  For 
the  25  measurements  we  then  made,  we  found  the  Tran-Pecos  specimens  fell 
within  the  range  of  the  25  S,  marnocfyi  measurements. 

We  have  measured  mature  S.  marnoclyt  from  22.55  mm.  to  33  mm.  and  two 
28  mm.  S.  gaigeae.  We  find  our  measurements  are  recorded  thus:  Width  of 
head  in  body  length  2.66-2.8  in  the  two  S.  gaigeae  specimens  within  the  whole 
range  of  2.66-3.29  in  our  S.  marnocfyt  specimens.  Or  put  as  the  dcscribcr  docs 
(ratio  of  width  of  head  to  body  length)  it  would  be  0.357-0.375  for  S.  gaigeae 
and  0.30-0.375  for  S.  marnoctyi.  The  actual  measurements  for  the  width  of 
head  in  the  two  S.  gaigeae  arc  10.5  mm.  and  ro  mm. 

Marnock's  Frog 

Syrrhophus  marnoc\ii  Cope.  Plate  LXXXIV;  Map  25. 

Range:  Helotes  near  San  Antonio,  San  Marcos,  and  Austin,  to  San  Angclo, 
Tex.,  and  Sonora. 
Habitat:  Cracks,  crevices,  caves  in  limestone  ledges  of  the  hills  and  ravines. 


396  HANDBOOK  OF  FROGS  AND  TOADS 

We  made  our  captures  under  large  flat  stones  near  limestone  ledges,  one  above 
the  wall  of  a  ravine. 

Size:  Adults,  7<6-i%  inches  (22.0-39.5  mm.). 

Our  two  specimens  were  22.5,  26  mm.  (See  S.  camfi  for  detailed  measure- 
ments.) 

The  American  Museum  has  one  (no.  22664)  collected  by  Byron  C.  Marshall 
at  Ezell's  Cave,  San  Marcos,  Tex.,  May  12, 1928,  which  is  24  mm. 

The  Museum  of  Comparative  Zoology  has  two  specimens  (nos.  18527, 
185273)  from  Hclotes,  Tex.  They  measure  22  and  25  mm. 

The  Baylor  University  Museum  has  G.  W.  Marnock's  collection,  some 
adults  of  which  vary  from  23-55  mm.,  and  others  taken  by  W.  J.  Williams  in 
1927  and  1928.  One  of  these  is  31  mm. 

General  appearance:  This  small,  flattened  frog  has  a  long  flat  head,  with  a 
broad  space  between  the  eyes,  which  are  large  and  prominent,  indicating  noc- 
turnal activity.  In  general  color,  the  back  is  greenish,  spotted  with  brown,  the 
under  parts  light-brownish  vinaceous,  the  legs  crossbarred.  The  nostril  is  far 
in  front  of  the  eyes,  which  are  as  far  apart  as  the  breadth  of  the  hump.  The 
skin  is  smooth.  There  aie  one  or  two  white  or  deep  colonial  bufl  tubercles  be- 
low and  behind  the  tympanum. 

Color:  Helotcs,  Tex.,  May  5, 1925.  General  color  greenish  or  olive  lake;  top 
of  head  from  forward  part  of  upper  eyelid  mignonette  green  or  grape  green  or 
light  yellowish  olive,  the  upper  eyelid  rainctte  green  and  unspotted.  In  fact 
whole  top  of  head  from  upper  eyelid  forward  is  unspotted  and  sharply  marked 
oil  by  the  vertical  purplish  lorcal  area.  The  dorsum  from  upper  eyelids  back- 
ward becomes  ecru-olive  and  the  upper  parts  of  forehmbs  arc  the  same  or 
lighter,  a  deep  colonial  bufT.  On  the  upper  surface  of  hind  limbs  this  color 
changes  to  olivc-ocher,  yellow-ocher,  old  gold,  or  yellower  than  any  of  these 
three.  Spots  on  back,  black,  sorghum  brown,  or  vinaceous-brown.  The  tympa- 
num and  across  shoulder  insertion  is  light  purplish  vmaccous.  The  groin  is 
purplish  vinaccous.  Underside  of  hind  legs,  forelegs,  and  across  breast  arc  light 
brownish  vinaccous.  Throat  pale  grayish  vmaceous;  sides  light  brownish 
vinaccous  or  pale  grayish  vinaceous;  back  of  eye,  some  of  tympanum,  canthus 
extending  around  snout,  labial  rims,  sorghum  brown;  chin  specked  with 
sorghum  brown.  On  sides  of  body  are  some  white  or  pallid  brownish  drab  or 
pale  russtan  blue  spots;  one  or  two  white  or  deep  colonial  buff  or  chamois 
tubercles  below  and  behind  the  tympanum.  Belly,  pale  yellow-green  or  pale 
fluorite  green;  a  purplish  vinaccous  blood  vessel  down  middle  of  belly  divides 
the  color  into  two  parts.  Iris  dull  green-yellow  above  and  below  with  sorghum 
brown  vermiculatums  or  spots  on  it,  or  upper  part  \\ith  one  wavy  longitudinal 
line  and  then  small  spots;  front  and  rear  of  pupil  dark  vinaccous-brown,  thus 
from  snout  to  rear  of  eye  one  continuous  vitta. 

In  some  alcoholic  specimens,  the  upper  parts  from  head  to  vent  are  very 
finely  spotted;  and  in  some  of  the  smaller  specimens,  the  following  transverse 


ROBBER  FROGS:  LEPTODACTYLIDAE  397 

bars  on  the  hind  legs  may  be  indicated,  two  on  the  tarsus,  two  on  the  tibia,  one 
just  below  the  knee,  and  two  on  the  femur.  On  the  tibia  and  femur,  one  of  the 
two  bars  may  be  light-centered. 

Structure:  Small  flattened  body;  eyes  prominent,  widely  separated  by  broad 
flat  space;  fingers  and  toes  long  and  slender  with  prominent  tubercles,  which  if 
viewed  from  the  side  give  the  digits  a  prominent  saw-toothed  appearance;  tips 
of  fingers  are  expanded  and  truncate,  toes  less  so.  Wrist  of  forehmb  extends 
beyond  extremity  of  muzzle.  Foot  10-11  mm.  Tarsus  6.5  mm.  No  vomcrine 
teeth. 

In  several  we  have  seen  white  tubercles  on  right  and  left  sides  of  lower  belly, 
also  on  rear  of  femur.  Arc  they  natural  or  parasites? 

Voice:  This  is  a  cricket! ike  chirp  of  one  or  two  notes  of  an  instant's  duration, 
possibly  followed  by  a  trill  of  two  or  three  notes. 

May  6,  1925,  Helotes,  Tex.  While  I  was  describing  this  specimen  it  chirped 
on  three  different  occasions,  and  reminded  me  of  the  chirping  S.  cam  pi  of  a 
week  or  more  ago  at  Brownsville,  Texas. 

We  searched  with  flashlights  more  than  once  along  the  muddy  bank  of 
Helotes  Creek  at  the  flat  crossing  near  Marnock's  home  for  the  source  of  a 
similar  chirp.  We  thought  it  might  belong  to  a  frog,  but  search  as  we  would, 
we  could  not  find  the  musician.  It  was  back  a  little  from  the  edge  of  the  water, 
where  the  mud  was  soft,  punched  by  cows,  and  more  or  less  covered  with 
grassy  tussocks.  Similarly  we  searched  in  Brownsville  on  Camp  Brown 
grounds  for  a  chirp  on  the  side  of  a  grassy  ditch.  We  had  no  success. 

May  10,  Helotes,  Tc\.  On  our  hill,  we  went  walking  at  6  P.NI.  Nighthawks 
booming;  a  few  robber  frogs  calling.  Below  limestone  ledge  heard  a  crickctlikc 
call  frequently  followed  by  a  trill  of  two-three  notes.  A  froglikc  note,  Acru- 
like  only  shorter.  Is  it  Marnock's,  in  .imong  rocks  and  oaks? 

Breeding:  Development  is  probably  within  the  egg  and  during  April  and 
May. 

A  few  of  the  larger  specimens  in  collections  taken  in  May  look  to  be  ripe, 
and  almost  all  the  specimens  captured  have  been  secured  in  that  month.  We 
suspect  that  the  month  of  May  is  an  important  part  of  the  breeding  period. 
Eggs  not  described. 

It  seems  likely  that  these  frogs  hatch  fully  developed  from  the  egg,  just  as 
Dr.  Stejnegcr  and  Dr.  Thomas  Barbour  suggest  likely  for  S.  cumpi. 

There  is  probably  no  free-swimming  tadpole  and  no  transformation  in  that 
sense.  There  is  no  record  of  a  specimen  of  the  small  si/,c  (5  or  6  mm.)  of 
S.  campt.  The  smallest  we  have  seen  are  two  taken  by  W.  ].  Williams.  They 
are:  Baylor  University  Museum  no.  2185  taken  at  San  Marcos,  March  },  1927, 
and  another,  no.  3853,  taken  at  San  Marcos,  June  1928.  Each  is  17  mm.  in  length 
of  body. 

Journal  notes:  May  5,  1925,  Helotes,  Tex.  In  Helotes  Creek  Canyon  above 
camp  .  .  .  where  a  side  ravine  comes  in,  we  were  searching  for  a  poinsctt's 


398  HANDBOOK  OF  FROGS  AND  TOADS 

lizard  we  had  seen.  .  .  .  About  3-4  feet  farther  up  the  side  of  the  canyon  was 
a  large,  flat,  loose  stone  on  a  stony  slope.  I  lifted  it  and  a  small  frog  hopped 
out.  I  got  the  impression  of  a  greenish  frog.  I  dropped  the  stone  quickly  to  see 
where  the  leaping  frog  went  but  lost  it.  Then  I  lifted  the  stone  again,  and  at 
first  my  eye  did  not  espy  the  second  frog.  Had  it  been  a  Camp's  frog  it  would 
have  leaped.  Presently  my  eye  saw  a  Syrrhophus.  It  is  Marnock's  frog.  It  is 
bigger  than  Camp's,  but  much  like  it. 

Our  second  frog  was  caught  May  10  on  Marnock's  hill,  back  of  the  store  at 
4  P.M.  In  ihe  morning  it  rained  hard  until  i  P.M.;  then  we  went  to  Marnock's 
hill.  East  of  the  store  one-half  mile,  found  on  top  of  the  hill  a  pebbly  place 
with  a  few  flat  stones.  A  banded  gecko  and  Marnock's  frog  were  under  a  stone. 
It  was  moist,  black  dirt  beneath  the  stone,  and  it  was  in  a  more  or  less  open 
place  with  dwarf  oaks  about. 

June  23, 1930,  San  Antonio.  I  left  at  8  o'clock  for  Helotes  with  Gable  and  his 
son,  Hugh.  They  parked  their  car  at  a  crossing  above  G.  W.  Marnock's  house. 
They  started  up  a  horseshoe  hill  I  never  was  on  in  1917  or  1925. 1  suspect  this 
was  Marnock's  special  hill  for  collecting.  His  house  is  in  the  opening  of  the 
horseshoe.  After  we  reached  the  top,  we  went  down  toward  a  glen  where  there 
had  been  seepage  earlier.  In  this  place  they  had  found  two  Marnock's  frogs, 
which  they  gave  to  Mr.  Parks  to  send  to  General  Biological  Supply.  One  of 
these  very  frogs  we  received  this  spring.  Their  habitat  was  much  the  same 
as  that  of  those  found  by  us. 

Authorities9  corner: 
W.  J.  Williams,  Tex.,  1927,  p.  7. 
}.  C.  Marr,  Gen.,  1944,  p.  480. 


FAMILY  RANIDAE 

Genus  RAN  A  Linne 
Maps  26-36 

Texas  Gopher  Frog,  Southern  Crayfish  Frog,  Southern  Gopher  Frog 

Rana  areolata  areolata  Baird  and  Girard.  Plate  LXXXV;  Map  26, 

Range:  Matagorda  Co.,  Tex.,  north  to  McCurtain  Co.,  Oklahoma,  and  La- 
fayette Co.,  Ark.;  probably  also  in  extreme  northwestern  Louisiana  (L.  Stej- 
neger  and  T.  Barbour,  Check  List,  1943,  p.  53). 

Habitat:  "I  saw  but  one  specimen  of  this  frog  and  collected  that  on  June 
1 8  at  the  Machine  Gun  Range,  twelve  miles  out  from  Camp  |  Houston,  Tex.  |. 
It  was  a  half-grown  frog  and  was  hiding  Under  a  log  at  the  edge  of  a  pool" 
(P.  H.  Pope,  Tex.,  1919,  pp.  97-98). 

Size:  Medium,  length  21/2-?H  inches  (62-90  mm.). 

General  appearance:  This  is  a  stocky,  medium  to  large,  rather  rough- 
skinned,  brownish  frog  with  round  dark  spots  on  back  and  sides,  with  reticula- 
tions of  dotted  light  lines  and  with  prominent  dorsolateral  folds. 

"Rana  areolata.  .  .  .  Appears  to  be  closely  allied  to/?,  halecina  \pipiens\  and 
utricularia,  but  the  head  is  larger  and  the  spots  of  the  back  are  smaller  and  more 
numerous.  Male  with  an  external  vocal  vesicle  on  each  side  behind  the  angle 
of  the  mouth  .  .  ."  (G.  A.  Boulenger,  Gen.,  1882,  p.  41). 

According  to  Cope  (Gen.,  1889,  p.  409),  R.  v.  areolata  has  "length  of  head 
to  posterior  tympana  three  times  in  total;  tympanic  disk  round;  dorsal  spots 
well  separated;  nostril  equidistant  between  end  of  muzzle  and  eye,"  whereas 
R,.  v.  circulosa  has  "length  of  head  one  third  of  total;  tympanic  disk  variable, 
dorsal  spots  so  large  as  to  leave  only  circles  of  the  light  ground-color;  nostril 
nearer  eye  than  end  of  muzzle  in  young." 

Coin  and  Netting  (Fla.,  1940,  p.  146)  give  these  contrasting  characters. 
"Head  U-shaped  in  outline  when  viewed  from  above;  dorsum  often  smooth, 
or  nearly  so;  tibia  length  less  than  40  mm.  in  adults;  post-tympanic  fold  poorly 
developed;  dorso-lateral  folds  narrow  or  only  slightly  raised,  or  both.  .  .  . 
Rana  areolata  areolata  Baird  and  Girard.  Head  orbiculate  in  outline  when 
viewed  above,  dorsum  rugose,  tibia  length  more  than  40  mm.  in  adults;  post- 
tymphanic  fold  well  developed;  dorsolateral  folds  prominent.  .  .  .  Rana 
areolata  circulosa  Rice  and  Davis." 


400  HANDBOOK  OF  FROGS  AND  TOADS 

Bragg  (Okla.,  19423,  p.  18)  of  Oklahoma,  which  state  has  both  areolata  and 
circulosa,  characterizes  them  as  follows:  "Dorsal  surface  smooth;  dorsolateral 
folds  narrow  and  usually  not  well  developed;  legs  fairly  short  (tibiae  40  mm. 
or  less).  Distribution,  southeastern  Oklahoma.  Rana  areolata  areolata,  South- 
ern Crayfish  Frog.  Dorsal  surface  often  rugose;  dorsolateral  folds  well  de- 
veloped; leg  rather  long  (tibiae  more  than  40  mm.).  Distribution  northeastern 
Oklahoma.  Rana  areolata  circulosa  Rice  and  Davis  (Northern  Crayfish 
Frog)." 

Color:  Houston,  Tex.,  from  R.  H.  Vines  through  the  kindness  of  A.  C. 
Chandler,  March  5,  1948.  Three  males.  The  background  of  the  backs  ranges 
from  chalcedony  yellow  to  olive-yellow.  In  the  darkest  one  there  are  more 
speckles  on  the  background,  which  is  deep  olive-buff.  In  two,  the  heads  are 
much  darker  than  the  backs,  and  the  larger  spots  are  inconspicuous  there.  In 
one,  the  head  is  like  the  back,  the  spots  clearly  showing.  The  large  collapsed 
vocal  sacs  are  courge  green  to  light-hellebore  green.  When  one  frog  blew  up 
his  pouch  to  a  diameter  of  %  inch,  the  pattern  of  spots  showed  in  a  dull 
and  dusky  manner.  The  concealed  surfaces  of  hind  legs  and  the  groin  are 
suffused  with  citron  yellow  to  strontian  yellow. 

In  the  smallest  and  lightest  frog,  the  background  of  back  and  head  is  dark 
olive  buff  except  the  snout,  which  is  darker.  On  the  back  between  dorsolateral 
folds  are  three  irregular  rows  of  spots  of  buffy  olive,  outlined  by  primrose 
yellow.  On  the  side  are  three  more  irregular  rows,  but  these  spots  are  not  com- 
pletely encircled  with  light  and  hence  are  not  so  conspicuous.  They  are  light 
brownish  olive  on  a  background  of  vinaceous-buff  to  wood  brown,  which  is 
marked  with  conspicuous  round  spots  of  ivory  yellow.  The  center  of  the 
tympanum  is  marguerite  yellow,  ivory  yellow,  or  white.  The  bars  on  the  legs 
are  very  prominent,  3  on  tibia  and  2  on  thigh  with  i  at  the  knee,  and  are  col- 
ored and  outlined  like  the  spots  of  the  back.  On  the  tibia  each  interspace  be- 
tween bars  bears  a  thin  line  of  light  brownish  olive.  On  tarsus  and  outer  edge 
of  foot  are  2  bars.  The  exposed  surface  of  the  femur  is  very  spotted  but  the 
spots  are  not  outlined  and  their  color  is  olive-yellow.  This  spotted  pattern  is 
revealed  on  the  venter  and  on  the  rear  margin  of  the  femur.  The  ventral  sur- 
face of  the  femur  is  light  vinaceous-fawn.  The  forehmbs  are  not  conspicuously 
barred,  being  only  irregularly  spotted.  The  under  surface  is  white.  The  edge 
of  upper  jaw  from  eye  backward  is  mottled,  and  the  rim  of  lower  jaw  bears  £ 
few  dark  spots.  There  is  a  spot  on  each  eyelid  and  one  on  the  interorbital  space. 
The  eye  is  large  and  bulging,  the  black  iris  bearing  fine  dots  of  zinc  orange 
with  the  warm  buff  pupil  rim  notched  below. 

In  the  largest  frog,  the  light  spots  of  the  sides  are  often  centered  with  pin 
points  of  black  and  are  raised  warts  on  mounds.  Similar  wartlike  spots  cross 
the  rear  of  the  sacral  region.  The  black  pin  points  are  so  conspicuous  on  the 
back  that  this  is  a  very  speckled  frog.  Between  the  coastal  folds  are  5-7  low 
irregular  broken  rows  of  longitudinal  folds. 


FROGS:  RAN1DAE  401 

Structure:  Length  of  head  contained  in  total  length  three  times.  Length 
of  leg  to  heel  equals  length  to  eye  or  nostril.  Tibia  longer  than  femur. 

"Rana  areolata,  B.  &  G.  Head  very  large,  sub-elliptical;  snout  prominent, 
nostrils  situated  halfway  between  its  tips  and  the  anterior  rim  of  the  eyes, 
which  are  proportionally  large.  The  tympanum  is  spherical,  and  of  medium 
size;  its  central  portion  is  yellowish-white,  whilst  its  periphery  is  black.  The 
body  is  rather  short  and  stout;  the  limbs  well  developed;  the  fingers  and  toes 
very  long  without  being  slender.  The  ground  color  of  the  body  and  head  is 
yellowish-green,  marked  with  dark  brown.  Besides  there  arc  from  thirty  to 
fifty  brown  areolae;  margined  with  a  yellowish  line.  The  upper  part  of  the 
limbs  is  of  the  same  color  as  the  body,  but  instead  of  areolae,  transverse  bands 
of  brown  are  seen  on  the  hind  ones.  The  lower  part  of  the  head  and  body  is 
yellowish,  with  small  dusky  spots  along  the  margin  of  the  lower  jaw,  and 
under  the  neck.  A  specimen  three  inches  and  a  half  long  was  found  at  India- 
nola  [Texas],  and  a  very  small  one  on  the  Rio  San  Pedro  of  the  Gila"  (S.  F. 
Baird  and  C.  Girard,  Gen.,  1852,  p.  173).  This  last  frog  might  well  be  another 
form,  or  did  R.  areolata  occur  in  Arizona?  (Note  suggestion  on  p.  516.) 

"Skin  rough,  with  elongated  warts  on  back  and  sides.  Lateral  folds  con- 
spicuous. Tibia  ridged  lengthwise.  No  glandular  fold  along  the  jaw.  A  distinct 
fold  over  the  ear  from  eye  to  shoulder.  Under  and  posterior  surface  of  femur 
granulated.  Head  large;  unusually  thick  through.  Muzzle  long;  space  between 
eyes  greater  than  width  of  eyelid;  nostrils  nearer  to  the  end  of  the  muzzle 
than  to  the  eye;  eyes  large;  ear  half  to  two-thirds  size  of  eye.  Foot  with  a  tarsal 
fold;  webs  short  (three  joints  of  fourth  toe  free);  inner  sole  tubercle  small, 
no  outer  tubercle,  tubercles  under  toe-joints  prominent"  (M.  G.  Dickerson, 
Gen.,  1906,  pp.  192-193). 

Breeding:  February  to  June. 

"As  regards  an  easy  way  of  capturing  Texas  gopher  frogs,  there  is  no  such 
thing  as  an  easy  way  in  my  limited  experience  with  the  species.  In  fact,  I  cap- 
tured so  few  of  them  and  those  under  conditions  of  unusual  expenditure  of 
energy  that  even  at  this  date  I  can  remember  each  individual  capture.  I  have 
verified  my  memory  by  looking  up  my  notes. 

"The  first  is  a  male  captured  under  circumstances  which  suggested  the  mat- 
ing urge  was  upon  him.  The  second  was  observed  while  feeding  on  insects 
under  a  street  light  on  Main  Street,  just  outside  Hermann  Park.  It  was  cap- 
tured after  a  merry  chase  that  left  me  rather  muddy.  I  was  returning  to  the 
Institute  when  Dr.  and  Mrs,  Chandler  stopped  their  automobile  by  the  curb 
for  a  little  conversation.  The  remaining  frogs  were  all  captured  on  February 
ii,  1930,  after  a  careful  stalk  of  a  breeding  pool  where  the  frogs  were  singing. 
In  this  case  the  stalk  was  performed  so  slowly  that  more  than  an  hour  was  cov- 
ered in  approaching  the  pool  where  three  clasping  pairs  were  taken.  The  fe- 
males extruded  their  eggs  that  night.  Incidentally  these  frogs  secreted  a  foamy, 
sour-smelling  material  that  made  my  hands  itch. 


402  HANDBOOK  OF  FROGS  AND  TOADS 

"I  think  the  only  means  of  capturing  these  frogs  with  any  assurance  of  suc- 
cess would  be  to  stalk  the  animals  in  the  breeding  pools.  I  made  many  unsuc- 
cessful attempts  to  capture  these  frogs  before  I  approached  sufficiently  slowly 
and  carefully  to  get  them  under  the  flash  light.  On  land  these  frogs  are  very 
active.  My  experience  suggesting  they  are  even  more  difficult  to  capture  on 
land  than  the  southern  leopard  frog,  but  in  the  water  they  are  very  slow  to 
move  and  can  be  seized  easily.  However,  they  are  very  shy  and  if  one  ap- 
proaches the  breeding  pool  at  a  normal  pace,  these  frogs  are  nowhere  to  be 
seen  when  one  arrives. 

"The  pool  where  I  caught  the  mating  pairs  was  located  northwest  of  the 
chemistry  building,  but  all  that  portion  of  the  campus  is  now  in  cultivation 
and  I  dare  say  that  the  pool  has  been  destroyed;  the  frogs  dispersed  to  other 
areas.  Breeding  pools  are  easily  located  by  the  mating  song,  which  is  produced 
after  each  heavy  rain  in  late  January  and  February.  The  mating  song  is  easily 
learned,  because  so  far  as  I  could  tell  it  is  exactly  like  the  mating  song  of  the 
most  common  toad  of  Houston,  namely,  Bufo  valhceps.  However,  the  toad 
sings  in  May  and  June,  while  the  gopher  frog  sings  in  the  winter"  (letter  from 
P.  D.  Harwood  to  A.  C.  Chandler,  June  12,  1946). 

Journal  notes:  Northeast  of  Edinburg,  Tex.,  Aug.  3,  1942.  Ten  or  fifteen 
miles  north  of  Raymonclville  a  dead  Rana  pipiens  or  areolata  in  the  road. 
Mesquite  on  either  side.  A  many-spotted  frog  we  could  see,  and  little  else. 
Another  frog  so  crushed  I  could  not  tell  it. 

Authorities*  corner: 
M.  C.  Dickerson,  Gen.,  1906,  p.  193. 
C.  E.  Burt  and  M.  D.  Burt,  Tex.,  19293,  p.  6. 
A.  N.  Bragg  and  C.  C.  Smith,  Okla.,  1943,  p.  107. 

Northern  Gopher  Frog,  Gopher  Frog,  Crayfish  Frog,  Hoosier  Frog, 
Crawfish  Frog,  Ring  Frog 

Rana  areolata  ctrculosa  Davis.  Plate  LXXXVI ;  Map  26. 

Range:  "From  Rogers  and  Tulsa  counties,  Oklahoma,  north  through  east- 
ern Kansas,  eastward  across  central  Missouri  and  Illinois  to  Bcnton  and  Mon- 
roe counties,  Indiana  (possibly  to  Greene  county,  Ohio),  and  southward  in  the 
Mississippi  Valley  through  western  Kentucky  and  Tennessee  to  Pontotoc4 
County,  Mississippi"  (L.  Stejacger  and  T.  Barbour,  Check  List,  1943,  P-  54)- 

Habitat:  Old  burrows,  commonly  crayfish  burrows  in  the  vicinity  of  ponds. 

In  1910,  "Professor  La  Rue  found  the  frogs  in  the  mammal  burrows  along 
the  shores  of  the  ponds,  as  well  as  in  crayfish  holes,  but  it  is  probable  that  they 
were  only  temporarily  occupying  the  former  during  the  spawning  season  for 
we  were  unable  to  discover  any  mammal  burrows,  either  in  the  vicinity  of 
ponds  or  elsewhere  inhabited  by  frogs"  (C.  Thompson,  111.,  1915,  p.  6.)  "The 
old  burrows  occupied  by  crayfish  were  entirely  without  chimneys,  and  were 


FROGS:  RANIDAE 


4°3 


Plate  LXXXV.  Rana  areolata  areolata  Plate  LXXXVl.  Kana  areowiu  t»t»^». 

tyu\  Maies  1,4.  Females  (X]/6)-  2.  Female  (X%)-  3- 

VA  Male 


4o4 


HANDBOOK  OF  FROGS  AND  TOADS 


approximately  round  at  the  entrance,  which  had  a  diameter  of  about  three 
inches.  The  entrance  was  more  or  less  overhung  with  grass,  and  at  one  side 
was  a  small  bare  space  about  six  inches  in  diameter.  .  .  .  The  burrows  oc- 
cupied by  frogs  differed  but  slightly  from  those  just  described"  (same,  p.  3). 

"The  habitat  is,  in  general,  low  meadowland,  which  is  sufficiently  moist  to 
harbor  crayfishes.  The  burrows  of  the  latter  are  necessary,  apparently,  for  the 
well-being  of  this  frog,  which  lives  in  them  during  the  day.  .  .  .  Holes  with 
funnels,  of  course,  can  be  inhabited  by  nothing  but  crayfish;  but  frequently  or 


Map  26 

usually  they  have  nothing  but  the  smooth  platform  in  front — always  in  this 
case  they  are  inhabited  by  frogs.  These  platforms  serve  to  prevent  grass  from 
growing  at  the  entrance  and  provide  mounds  upon  which  the  frogs  sit  at  night 
or  early  in  the  morning  to  cjtch  stray  insects"  (H.  M.  Smith,  Kans.,  1934,  pp. 
480-481). 

"During  the  postbrcedmg  season  ctrcttlosa  resorts  to  crawfish  burrows  and, 
therefore,  is  not  found  in  streams  or  ponds.  Dr.  Wilfred  Crabb  and  Mr.  George 
Wiseman  collected  four  specimens  on  May  6,  1942  at  Stockport,  Van  Buren 
County.  The  following  habitat  description  is  quoted  from  their  field  notes: 
'The  frogs  were  found  sitting  near  the  openings  of  holes  into  which  they  re- 
treated when  disturbed.  They  were  extricated  by  fish  hooks  attached  to  the 
ends  of  slender  sticks.  The  holes  were  apparently  vacated  crayfish  burrows. 


FROGS:  RANIDAE 


405 


One  was  three  feet  deep  with  18  inches  of  water  in  the  bottom,  another  was  28 
inches  with  14  inches  of  water.  Two  frogs  were  usually  present  in  the  same 
burrow  although  the  large  one  [adult  male;  snout  to  vent  length  67  mm.]  was 
alone.  There  were  about  12  holes  within  a  radius  of  50  feet  and  all  contained 
frogs.  They  were  located  on  a  short-grazed  bluegrass  pasture  with  a  slope  of 
7  per  cent.  The  day  was  cloudy  and  cool  (about  40°?.)  with  a  fresh  west  wind.' 
Another  specimen  taken  in  a  similar  situation  the  same  day  in  an  adjoining 
section  was  an  adult  female  101  mm.  in  length"  (R.  M.  Bailey,  la.,  1943,  p.  350). 

Size:  Adults,  2^-4  %  inches.  Males,  63-104  mm.  Females,  75-11?  mm. 
Among  Percy  Viosca,  Jr.'s  material  from  Ottawa,  Kansas,  from  Gloyd,  were 
males  105,  108,  m  mm.  and  one  92  mm.,  having  thumb  and  sacs  well  de- 
veloped. At  Monroe,  La.,  P.  Viosca,  Jr.,  and  H.  B.  Chase  had  caught  a  female 
of  87  mm.  (maturity  not  certain)  and  one  young  46.5  mm. 

General  appearance:  This  is  a  large  brownish  frog  with  round  dark  brown 
spots  on  back  and  sides.  These  spots  may  vary  in  size,  as  large  or  small.  They 
are  in  three  or  four  rows  between  and  sometimes  extending  onto  the  dorso- 
lateral  folds  and  several  irregular  rows  or  groups  on  the  sides.  The  spots  are 
surrounded  with  grayish  white  borders,  which  on  the  lower  sides  become  the 
background  color  between  the  spots.  The  skin  of  the  back  is  roughened  with 
tubercles.  There  are  one  or  two  long  glandular  folds  on  the  tibia.  There  is  con- 
siderable greenish  yellow  in  the  groin  and  on  the  concealed  portions  of  legs 
and  feet.  There  is  a  prominent  dark  bar  on  the  brachmm,  a  broad,  fleshy, 
spotted  band  along  the  jaw,  and  a  light  center  to  the  tympanum.  The  light 
color  on  the  arms  and  jaw  is  grayish.  There  are  several  smaller,  hglu-rimmccl, 
dark  spots  on  eyelids,  between  the  eyes  and  on  top  of  the  snout.  The  legs  aic 
prominently  barred  with  brown  and  cream  or  gray.  In  this  group,  the  frogs 
have  a  few  dark  spots  just  forward  of  the  arm  insertion.  When  cold  and  wet, 
the  frogs  were  very  dark.  The  vocal  sacs  of  the  males  are  large  and  conspicuous 
when  collapsed,  and  bluish  gray-green  in  color.  When  expanded  they  form 
large  balls  on  either  side  above  the  arms.  (When  plowed  out  in  early  spring 
they  are  so  dark  as  to  be  almost  blackish.) 

Color:  Calhoun,  111.,  from  CI.  A.  Nicholas,  Feb.  27,  iy$o.  Mule.  Down  the 
back  between  dorsolatcral  folds  arc  three  rows  of  spots.  Back  of  the  hump 
mainly  two  rows.  Spots  mummy  brown,  raw  umber,  brownish  olive,  or  clove 
brown.  These  are  encircled  by  thin  lines  of  ivory  yellow  or  cartridge  buff  or 
marguerite  yellow.  This  color  merges  into  interstices  between  spots.  These 
interstices  are  covered  with  small  dots  of  the  four  browns  given  above  for  big 
spots,  producing  in  effect  a  light  grayish  olive  or  grayish  olive.  This  color  on 
top  of  forearm  becomes  light  mineral  gray.  It  is  grayish  olive  on  hind  legs. 
The  femur,  tibia,  and  tarsus  have  mummy  brown  or  one  of  the  other  three 
browns  in  their  conspicuous  crossbars.  The  interspaces  between  bars  are  like 
interstices  of  back,  but  with  small  partial  crossbars  of  the  darker  color.  On 
sides  are  at  least  two  rows  of  large  spots;  mummy  brown  bar  on  forearm  in- 


406  HANDBOOK  OF  FROGS  AND  TOADS 

sertion.  Underside  of  forefeet  and  hind  feet  pale  vinaceous-drab  or  light  gray- 
ish vinaceous.  Under  parts,  throat,  breast,  belly,  and  lower  sides  pale  cinnamon- 
pink  or  cartridge  bufl  (before  molting  it  was  colonial  buff  or  deep  colonial 
bufl) .  Same  color  on  fore  half  of  venter  of  hind  legs  and  underside  of  fore- 
arms. Underside  of  femur,  down  middle,  pale  vinaceous-drab  or  light  grayish 
vinaceous.  Top  of  foot  and  tarsus  yellow  ocher  or  honey  yellow.  Wash  of  picric 
yellow  on  rear  of  arm,  axil,  groin,  front  of  thigh,  and  rear  of  tibia.  On  rear  of 
thighs  and  somewhat  in  front  oi  thighs  the  color  becomes  oil  yellow.  Edge  of 
lower  eyelid  bluish  gray-green  or  parula  blue.  Pupil  horizontal,  broken  in 
middle  below.  Over  pupil  obscure  longitudinal  band  of  xanthine  orange  in 
front  and  rear  or  with  considerable  empire  or  pinard  yellow.  Pupil  has  rim  of 
this  yellow.  Iris  in  general  black.  Vocal  sacs  tea  green  to  celandine  green. 

Female.  Encircling  light  lines  around  large  dorsal  spots,  general  interstices, 
interstices  of  hind  legs,  and  light  spot  in  tympanum,  pale  olive-buff  or  tilleul 
buff,  on  sides  becoming  even  vinaceous-buff.  The  dorsolateral  fold  is  tilleul 
buff  or  vinaccous-buff.  This  interstitial  color  is  with  many  fine  dots  of  olive 
brown  or  deep  olive.  The  large  dorsal  spots,  three  rows  forward  and  two  back- 
ward of  the  hump,  are  clove  brown  or  fuscous.  These  spots  are  smaller  on  sides 
and  become  natal  brown.  The  crossbars  on  hind  legs  and  spots  on  forelimbs 
are  like  spots  of  dorsum.  Spot  across  arm  insertion  in  front  and  one  just  beyond 
insertion  on  arm  in  axil.  Side  of  head,  edge  of  lower  jaw,  and  venter  ahead  of 
arm  insertion  with  small  olive-brown  or  clove  brown  spots,  giving  side  of  head 
a  speckled  appearance.  Axil,  rear  of  arm,  groin,  front  of  thigh,  and  rear  of 
thigh  greenish  yellow  or  green-yellow.  On  reticulated  rear  of  femur  this  be- 
comes oil  yellow  or  courge  green.  Dark  color  is  fuscous.  Front  of  tibia  and  top 
of  foot  cream-buff.  Under  parts  are  pale  pinkish  buff  or  marguerite  yellow. 
Thigh  on  underside  has  same  light  grayish  vinaceous  of  male.  Also  same  color 
on  underside  of  foot  and  hand.  Ins  black  or  fuscous.  Pupil  nm  vmaceous- 
tawny  or  vinaceous-fawn.  Over  pupil,  a  horizontal  band  or  dash  of  these 
vinaceous  colors.  Same  color  in  fine  dots  on  lower  iris  and  outer  part  of  upper 
iris. 

Structure:  Large;  skin  warty  on  back  and  sides;  head  shorter,  mouth  smaller, 
and  hind  limbs  longer  than  R.  capita;  males  having  prominent,  collapsed, 
pleated  vocal  sacs  resting  outside  like  folds  of  skin  and  continuing  along  the 
sides  as  folds  past  the  axil,  the  middle  of  the  sac  being  back  of  the  tympanym; 
thumb  somewhat  enlarged  in  the  male;  eye  conspicuous,  but  small  in  relation 
to  snout;  fourth  toe  very  long;  a  thickened  or  fleshy  band  along  the  edge  of  the 
jaw.  On  the  breast,  the  arm  insertion  with  the  pectoral  girdle  is  conspicuously 
indicated  by  a  triangle  of  much  thinner  skin,  the  base  at  arm's  insertion,  the 
point  at  the  pectoral  region;  waist  slightly  broader,  thus  making  whole  form 
less  wedge-shaped  than  R.  capita. 

In  Jordan's  Manual  of  Vertebrates  of  the  Northern  United  States  .  .  .  (2d 
ed.,  rev.  and  enl.,  Chicago,  187(8),  page  355,  we  have  this  description:  "R.  circti- 


FROGS:  RANIDAE  407 

losa,  Rice  and  Davis  (sp.  nov.)  Hoosier  Frog.  Head  broad;  body,  head  and 
sides  with  the  ground  color  largely  predominating,  and  with  narrow  rings  of 
a  greenish  slate  color,  which  becomes  larger  and  more  irregular  posteriorly; 
hind  legs  black,  crossed  with  irregular  lines  of  yellowish  slate  color;  fore  limbs 
similarly  marmorate;  tympanum  black  with  pale  ring;  below  chiefly  yellowish 
white;  toes  very  long;  size  medium,  L.  3/l>.  Benton  Co.,  Indiana,  lately  dis- 
covered by  Mr.  E.  F.  Shipman  (abridged  from  Mr.  Rice's  Notes)." 

"There  is  a  strong  dorsolateral  glandular  ridge  on  each  side,  and  between 
these  there  are  from  six  to  eight  narrow  glandular  folds  not  so  much  broken 
up  as  in  the  R.  a.  aesopus,  but  readily  becoming  indistinct  in  alcohol.  The  dorso- 
lateral fold  extends  nearly  to  the  groin.  Below  it  the  sides  are  crowded  with 
longitudinal  glandular  folds,  more  or  less  broken  up.  .  .  . 

"Since  the  above  was  written  I  have  been  able,  through  the  kindness  of  Pro- 
fessor Forbes,  of  the  university  at  Champaign,  111.,  to  examine  the  type  speci- 
men of  Messrs.  Rice  and  Davis.  It  differs  considerably  from  the  specimens 
above  described,  as  follows:  The  muzzle  is  not  protuberant,  so  that  the  nostril 
is  equidistant  between  the  end  of  the  muzzle  and  the  eye,  as  in  the  subspecies 
arcolata.  The  tympanic  disk  is  nearly  round,  and  its  long  diameter  is  three- 
fourths  that  of  the  eye.  This  specimen  has  twice  the  bulk.  In  other  respects 
it  does  not  differ.  A  very  strong  glandular  thickening  of  the  skin  extends  from 
the  eye  above  the  tympanum,  and  then  descends  posterior  to  it.  The  eyelid  also 
is  thickened. 

"Two  specimens  (No.  13828)  from  Olney,  III.,  also  received  since  the  above 
description  was  written,  explain  these  discrepancies.  The  larger  of  the  two 
agrees  with  the  type  in  all  respects,  but  the  smaller,  which  about  equals  the  type 
in  dimensions,  has  the  elongate  muzzle  of  the  small  ones  that  I  have  described 
above.  In  both  the  tympana  are  three-fourths  the  orbit,  and  in  neither  is  it  de- 
cidedly oval"  (E.  D.  Cope,  Gen.,  1889,  pp.  41  $-415). 

Voice:  "A  loud  trill,  hoarser  than  that  of  the  leopard  frog  and  pitched  some- 
what higher  than  that  of  Rana  catesbeinna"  (C.  Thompson,  Mich.,  icji5a,  p.  6). 
To  Mr.  Ackert,  the  call  sounds  "half -strangled"  as  if  it  had  its  mouth  half  out 
of  water. 

"The  call,  from  near  at  hand,  is  of  the  general  type  of  that  of  Rana  palustris, 
but  is  much  louder  and  quite  distinct  in  timbre.  It  may  best  be  described  as  a 
deep  guttural,  snoring  sound,  with  a  slight  upward  crescendo  at  the  end.  It 
is  given  at  intervals  of  perhaps  ten  to  fifteen  seconds,  though  we  did  not  time 
it"  (H.  P.  Wright  and  G.  S.  Myers,  Ind.,  1927,  p.  174). 

"The  song  of  Rana  arcolata  was  most  often  heard  after  dark  although  on 
one  occasion  several  were  singing  and  splashing  in  a  roadside  pond  about  an 
hour  after  sundown.  The  voice  of  these  frogs  does  not  have  the  prolonged 
resonance  of  that  of  the  bullfrog,  R.  catesbeiana,  although  it  is  almost  as  deep 
and  seems  to  have  even  more  carrying  power.  The  song  most  frequently  heard 
is  a  low-pitched,  drawn-out  guttural  note  which  may  be  suggested  by  the 


408  HANDBOOK  OF  FROGS  AND  TOADS 

syllables  "wurr-r-r-up"  accented  on  the  last.  It  is  repeated  several  times,  either 
from  the  surface  of  the  water  or  from  the  shore,  at  more  or  less  regular  inter- 
vals, varying  in  frequency.  The  vocal  sacs  of  the  males  are  lateral  and  rela- 
tively much  larger  than  those  of  R.  ptpicns.  When  singing  they  are  distended 
until  they  resemble  miniature  balloons,  each  one  almost  as  large  as  the  head 
itself"  (H.  K.  Gloyd,  Kan.,  1928,  pp.  117-118). 

"A  deeply  sonorous  and  resonant  'w-a-a-ah';  loud  and  of  exceptional  carry- 
ing quality.  From  mixed  choruses  this  stands  out  above  the  others;  clearly 
audible  for  over  half  a  mile.  The  strong  choruses  are  heard  in  April"  (R.  M. 
Bailey,  la.,  1944,  p.  17). 

Breeding:  March  to  mid-May.  The  National  Museum  (no.  48697)  has  a  ripe 
female  of  103  mm.  taken  at  Montgomery,  Miss.,  on  March  9,  1911,  by  Mr. 
Parker.  (See  Gloyd  in  "Authorities'  corner.")  We  ourselves  have  not  seen  the 
egg  masses,  but  they  are  doubtless  phnthhke  like  those  of  Rana  capita. 

On  April  1 1,  1926,  near  Bloomington,  IncL,  Herman  P.  Wright  and  George 
S.  Myers  found  17  egg  masses  of  this  species  in  a  small  pond  not  more  than 
90  feet  in  diameter. 

"I  have  found  these  frogs  singing  in  numerous  choruses  during  the  last  of 
March  and  first  of  April  m  southeastern  Kansas.  Near  Lawrence  they  were 
collected  while  breeding  on  April  27,  19.51.  In  1933  mated  pairs  were  collected 
at  the  same  locality  on  April  2r.  Gloyd  .  .  .  states  that  they  were  last  heard 
singing  in  1927  in  Franklin  county  on  April  9;  the  first  specimens  were  taken 
March  it.  ... 

"Temporary  pools  by  roadsides  and  in  pastures  are  chosen  in  which  to  breed 
and  lay  eggs.  Males  sing  at  the  edge  of  the  pools  or  out  in  the  water,  and,  al- 
though they  cease  singing  frequently  upon  the  approach  of  a  light,  yet  they 
will  remain  above  water  until  splashing  about  or  other  noises  cause  them  to 
cluck  beneath  the  water  or  to  sidle  back  into  their  holes.  In  some  pools  it  fre- 
quently was  possible  to  capture  them  after  they  disappeared  by  passing  the 
hands  back  and  forth  over  the  mud  and  grass  at  the  bottoms  of  pools,  where 
they  remain  hidden  until  further  disturbed  or  danger  is  past.  A  number  of 
specimens  placed  in  a  tank  with  three  or  four  inches  of  water  illustrate  well 
this  protective  instinct.  Upon  being  startled  by  some  sudden  activity  outside, 
they  duck  to  the  bottom  of  the  water,  close  their  eyes,  push  their  heads  down 
against  the  bottom  of  the  tank,  and  propel  themselves  blindly  forward  by  slow 
alternate  or  coincident  strokes  of  the  hind  legs,  holding  the  front  legs  against 
the  body.  Such  actions  in  pools  on  soft  earth  would  very  quickly  cover  them 
from  sight"  (H.  M.  Smith,  Kans.,  1934,  pp.  479-480). 

"Laid  in  large  plinth-like  masses  about  5-6  inches  in  diameter  in  shallow 
water  about  stems  of  grass,  etc.  Probably  about  7000  eggs  are  in  each  mass. 
Individual  eggs  are  rather  distinct.  The  outer  membrane  measures  about 
4.5-5.0  mm.  in  diameter,  the  inner  about  3.15,  and  the  vitellus  about  2.46-2.50. 
The  vitellus  is  considerably  larger  than  in  pipiens,  and  the  space  between  the 
inner  and  outer  membrane  is  greater  than  in  the  latter  species"  (same,  p.  479). 


FROGS:  RANIDAE  409 

The  transformation  time  is  the  first  week  of  July  (H.  P.  Wright  and  G.  S. 
Myers,  Ind.,  1927,  p.  174).  Size,  20-30  mm.  Herman  P.  Wright  and  George  S. 
Myers  were  to  have  described  the  tadpoles.  We  had  tadpoles  of  this  species 
from  Antioch  College,  Yellow  Springs,  but  they  were  without  eyes,  etc.,  i.e., 
experimental  specimens.  A  number  of  tadpoles  secured  4  miles  north  of  Bloom- 
ington,  Ind.,  April  12,  1940,  by  H.  T.  Gier  were  loaned  to  us  by  Mittleman. 
They  measured  39.5-51.0  mm.,  the  bodies  being  20-22  mm.  Most  of  them  were 
approaching  transformation.  Three  transformed  and  transforming  ones  were 
20,  20,  and  25.5  mm. 

The  Museum  of  Comparative  Zoology  has  two  specimens  (no.  1478),  col- 
lected in  Michigan  by  J.  G.  Shutc,  1863,  which  are  slightly  past  transformation. 
They  are  33  and  34.5  mm. 

"Sexual  maturity  is  apparently  attained  at  an  age  of  not  less  than  three  years. 
Three  early  spring  collections  of  overwintering  tadpoles  show  no  sign  of  ap- 
proaching metamorphosis,  but  since  a  unimodal  size  dispersion  is  evident  .  .  . 
[a  table  gives  overwintering  tadpoles  ranging  from  31-63  mm.],  transforma- 
tion may  be  assumed  to  occur  during  the  ensuing  summer.  Three  frogs  taken 
by  Crabb  and  Wiseman  on  May  6  were  immature;  they  have  snout  to  vent 
lengths  of  48  (  9  ),  49  (  9  ),  and  51  ( <J  )  mm.,  and  are  judged  to  be  two  years 
old,  having  transformed  during  the  preceding  summer.  The  adult  male  67 
mm.  in  length  taken  with  them  is  presumably  a  year  older;  it  is  possible, 
however,  that  he  transformed  early  in  the  summer  of  his  first  year  and  was 
mature  at  two  years  of  age.  Since  this  is  easily  the  smallest  adult  taken  it  is 
believed  the  bulk  of  breeding  specimens  are  at  least  four  years  old"  (R.  M. 
Bailey,  la.,  1943,  p.  350). 

Journal  notes:  Sept.  u,  1929.  At  n  A.M.  went  with  Dr.  Fernandus  Payne  to 
the  Rana  areolata  pond  found  near  Bloommgton,  Ind.,  in  1926  by  H.  P.  Wright 
and  G.  Myers.  It  is  the  only  pond  in  that  vicinity  in  which  they  have  found 
this  frog  breeding.  They  discovered  it  by  hearing  the  frog  chorus.  The  pond  is 
Vi  mile  from  the  road.  There  arc  cultivated  fields  (cornfields)  on  one  side 
of  it  and  meadow  on  another.  On  one  side  is  a  hedgerow  of  a  few  willows  and 
oaks,  and  near  by  are  a  few  shrubs,  but  in  the  main  the  pond  is  open.  It  was 
dry  in  the  summer  and  the  bottom  mud  cracked  into  blocks.  A  recent  rain 
formed  a  small  pool  in  the  center.  We  could  not  tell  where  m  the  surrounding 
country  the  Rana  a.  circulosa  would  be  likely  to  have  gone.  We  went  on  to 
Olney,  111.,  and  south  to  Calhoun  to  look  at  those  habitats.  There  we  looked 
up  Lee  Ackert,  who  went  out  as  a  young  lad  with  the  Thompson  girls,  when 
they  were  working  on  this  form.  He  took  us  to  their  farm  a  mile  or  so  east  of 
Calhoun.  Here  in  a  field  that  contained  an  open  pond  we  searched  for  the 
characteristic  holes  these  frogs  occupy.  He  said  the  holes  were  2V&-3  inches 
across.  We  found  nothing  promising  in  or  near  the  pond.  The  fields  were  so 
thickly  grown  up  to  grass  and  pasture  weeds  that  we  found  very  few  holes. 
The  boy  dug  up  two  or  three  but  had  no  luck.  He  said  he  has  plowed  them 
up  and  cut  them  in  plowing.  Thought  possibly  a  field  not  worked  this  year 


410  HANDBOOK  OF  FROGS  AND  TOADS 

would  be  better.  Went  to  a  pond  a  short  distance  north  and  showed  us  places 
there  where  the  "girls"  found  them. 

Mr.  Nicholas  reported  plowing  up  several  in  the  spring.  We  sought  him  i% 
miles  southeast  of  Calhoun.  He  took  us  down  his  lane  where  2  or  3  weeks  ago 
he  saw  a  Ran  a  a.  circulosa  at  the  entrance  of  a  2-3-inch  hole.  We  dug  and  dug; 
finally  at  about  4  feet  we  reached  water  level.  He  reached  in  several  times  and 
was  nipped  twice.  Proved  a  crayfish.  Did  he  drive  out  the  frog?  Went  to  house 
and  field  on  opposite  side  of  road.  In  a  lower  portion  near  woods,  which  he 
tells  us  is  covered  with  water  until  late  in  April,  we  searched  for  likely  holes. 
Found  one  but  it  was  filled  in.  Mr.  Nicholas  told  us  that  sometimes  two  or 
three  frogs  may  be  in  one  such  hole  with  heads  near  the  opening.  Sometimes 
they  appear  to  be  in  colonies  when  plowed  up.  They  appear  quite  blackish 
when  broken  out  and  look  almost  the  color  of  a  catfish.  They  work  down  into 
the  dirt  again  after  being  disturbed.  They  are  large,  heavy  frogs,  clumsy  and 
slow-moving.  His  place  is  near  the  source  of  Sugar  Creek.  Sugar  Creek  Prairie, 
a  known  habitat,  is  southwest  of  him  about  2^-3  miles  toward  Parkers- 
burg. 

During  three  warm  days  at  the  end  of  February,  1930,  he  caught  and  sent 
us  four  frogs,  one  male,  three  females,  from  this  prairie.  To  secure  one  of  these 
he  told  us  he  dug  down  5  feet. 

Authorities'  corner: 

G.  E.  Beyer,  La.,  1900,  p.  37.  J.  Hurter,  Mo.,  1911,  pp.  116-117. 

F.  A.  Hartman,  Kans.,  1907,  pp.  226-     H.  K.  Gloyd,  Kans.,   1928,  pp.   117- 

229.  1 1 8. 

Oregon  Red-legged  Frog,  Western  Wood  Frog 

Rana  aurora  aurora  (Baird  and  Girarcl).  Plate  LXXXVII;  Map  27. 

Range:  British  Columbia,  Washington,  Oregon,  south  along  the  coast  of 
California  to  Eureka. 

Habitat:  "Oregon  and  Washington  specimens  were  taken  among  the  ferns 
and  dense  vegetation  in  the  forests  of  the  coastal  belt"  (J.  R.  Slevin,  B.C.,  1928, 
p.  129). 

"This  frog  is  abundant  in  the  region  of  Portland  throughout  the  year  except 
in  December,  when  all  of  our  frogs  are  in  hibernation.  It  is  an  inhabitant  of 
the  deep  forests,  resting  under  the  cool  sword  ferns  of  this  region  during  the 
day  and  often  coming  forth  at  night  in  search  of  food"  (S.  G.  Jcwett,  Jr.,  Ore., 
1936,  p.  71). 

"It  is  usually  found  in  the  ferns  and  vegetation  of  the  woods,  or  in  nearby 
wet  areas"  (K.  Gordon,  Ore.,  1939,  p.  62). 

March  16,  1942,  Clam  Beach,  Calif.  In  a  pussy-willow  pond  took  an 
Amt>ystomci-l\ke.  mass  of  eggs.  In  a  pond  just  back  of  beach  dune  behind  a 
eucalyptus  cover  took  several  Ambystoma  masses  well  along  and  one  Rana 


FROGS:  RANIDAE 


411 


aurora  subsp.  Saw  two  Rana  aurora 
on  logs  and  one  gave  a  Rana  catlike 
call  when  it  leaped.  They  are  not  very 
fast  but  live  among  bad  tangles  of 
pussies  and  logs  and  tussocks. 

Size:  Adults,  r?4~3%  inches.  Males, 
44-63  mm.  Females,  52-87  mm.  Great- 
est size  less  than  Rana  aurora  dray- 
tonii.  Storer  (Calif.,  1925,  p.  231)  had 
males  50-58  mm.  and  females  57-76 
mm.  in  length. 

General  appearance:  Distinctly  a 
wood  frog,  the  mask  is  evident  in 
many.  It  is  medium  in  size,  moder- 
ately stout,  and  smooth-skinned.  The 
head  is  narrowly  oval  from  above,  the 
profile  thin.  The  back  is  brownish  or 
olive,  frequently  with  inky  spots. 
Sometimes  a  few  of  these  spots  have 
light  centers.  Frequently  there  is  a 
dark  bar  across  the  upper  arm.  There 
is  red  on  the  sides  of  the  body  and  on 
the  concealed  parts  of  the  legs,  feet, 
and  underarms.  The  light  line  along 
the  upper  jaw  ends  in  a  fold  at  the 
corner  of  the  mouth.  Groin  mottled. 

March  31,  1942.  J.  R.  Slater  com- 
pared three  Ranas  of  Oregon  and 
Washington. 

Mottled  red,  yellow,  and  green  in 
groin;  R.  aurora  aurora. 

Mottled  somewhat  in  groin  and 
lower  sides  yellowish,  cream;  R.  cas- 
cadac. 

Not  mottled  in  groin;  groin  red; 
R.  prcttosa. 

April  2,  Longview,  Wash.  Just  east 
of  Stella,  at  basaltic  falls  beside  road, 
next  the  water's  edge  took  one  small 
jR.  aurora  aurora.  At  west  edge  of  falls 
in  big  cavities  among  large  rocks  and 
logs  took  five  R.  aurora  aurora,  not  a 
fast  frog  nor  hard  to  catch,  yet  it  is 


R.  a.  c/rayfonii 
R.  cascadae 


Map  27 


4i2  HANDBOOK  OF  FROGS  AND  TOADS 

more  alert  than  R.  pretiosa  pretiosa.  The  groin  is  black  marked  with  con- 
spicuous spots  of  pale  greenish  yellow,  which  area  extends  slightly  onto  the 
belly. 

"The  Red-legged  frog  may  be  recognized  by  its  slender  form,  smooth  skin, 
dark  patch  behind  the  eye,  and  red  colour  on  sides  of  body  and  under  surfaces 
of  legs"  (G.  C.  Carl,  B.C.,  1943,  p.  48). 

Color:  Female.  Tacoma,  Wash.,  from  Prof.  James  Slater,  April  21,  1930. 
Underside  of  tibia,  front  half  of  venter  to  femur  and  along  sides  from  groin  to 
axilla  solid  coral  red,  jasper  red,  or  light  jasper  red.  A  wash  of  same  color  on 
hind  toes  and  membrane  of  web.  Top  of  back,  snout,  fore  limbs,  hind  limbs 
brownish  olive,  light  brownish  olive,  Saccardo's  umber  to  umber.  Sides  below 
costal  fold  with  considerable  black,  area  back  of  eye  more  or  less  black  but  not 
a  clear  vitta.  Deep  colonial  buff  or  deep  olive-buff  or  Naples  yellow  line  from 
shoulder  insertion  below  tympanum  to  below  eye  is  broken  up  on  side  of  snout 
with  many  black  dots.  Throat  heavily  specked  with  smoke  gray,  pale  olive- 
gray,  or  mouse  gray.  Breast  and  belly  white  but  with  much  gray  (of  the  shades 
listed  above)  on  it,  though  not  so  uniformly  overlaid  as  on  throat.  Groin  has 
black  and  some  light  dull  green-yellow  spots.  Eye  barium  yellow,  iris  border 
above  ochraceous-orange  or  zinc  orange  on  lower  ins  rim,  which  is  broken  in 
middle.  In  lower  iris  are  specks  of  same  color  as  lower  iris  border.  In  upper  ins 
is  a  little  ochraceous-orange.  Bars  on  hind  legs  not  very  distinct,  black  with 
many  ground  color  specks  on  them. 

Male.  Carbon  River  Valley,  Rainier  National  Park,  Wash.,  from  J.  R.  Slater, 
May  16,  17,  1930.  Dorsal  color  sulphinc  yellow,  citrine,  dull  citrine,  or  buffy 
citrine  with  few  faint  black  specks.  On  brachium  color  becomes  old  gold. 
Hind  legs  and  forelegs  same  color  as  dorsum.  Stripe  along  upper  jaw  to  over- 
arm insertion  barium  yellow  to  Naples  yellow  or  cream  color.  Mask  back  of 
eye  extends  diagonally  down  to  end  of  cream-colored  upper  labial  stripe.  Mask 
obscured  by  many  minute  oil  yellow  specks.  Iris  black  with  prominent  lemon 
yellow  or  light  greenish  yellow  bar  across  eye  above  pupil.  Lower  eye  mainly 
black  with  few  English  red  and  xanthine  orange  spots.  Upper  eye  with  few 
specks  above  light  greenish  yellow  bar.  On  sides  more  black  specks.  In  groin 
several  larger  viridine  green  spots.  Lower  sides,  undersides  of  tibia,  and  top  of 
hind  foot  and  femur  slightly  Etruscan  red,  old  rose,  Corinthian  red,  or  deep 
vinaceous.  Femur  hone  brown  with  white  specks.  Under  parts  dirty  white  or 
yellowish  glaucous.  In  general  venter  cloudy.  In  the  young  the  upper  labial 
light  line  is  particularly  conspicuous.  Iris  golden  yellow. 

Structure:  Frog  medium  in  size;  skin  smooth;  ridge  from  angle  of  eye  to 
shoulder  not  prominent;  dorsolateral  folds  indistinct;  webs  very  well  de- 
veloped, greater  in  males  than  females;  some  males  with  inconspicuous  groove 
across  middle  of  thumb  tubercle. 

"The  general  aspect  of  this  species  differs  greatly  from  all  its  congeners  in 
North  America.  The  length  of  the  body  and  head  together  is  three  inches  and 


FROGS:  RANIDAE  413 

a  half,  the  head  forming  nearly  one  third  of  this  length.  The  head  itself  is 
pyramidal,  pointed,  the  nostrils  situated  midway  between  the  anterior  rim  of 
the  eye  and  the  tip  of  the  snout.  Eyes  of  medium  size,  anterior  limbs  short; 
fingers  rather  long  and  slender.  The  body  is  orange  red,  with  here  and  there 
black  irregular  patches.  From  Puget  Sound"  (S.  F.  Baird  and  C.  Girard,  Ore., 
1852,  p.  174). 

Voice:  Seldom  recorded  in  the  field.  Like  the  wood  frog  it  has  a  short  breed- 
ing period  so  that  few  hear  it  to  know  it. 

Breeding:  These  frogs  breed  from  February  to  May.  Eggs  are  3.04  mm., 
outer  envelope  is  about  12  mm.  The  tadpoles  have  not  been  described.  They 
transform  at  11/io-%  inch  (17-21  mm,),  and  the  labial  teeth  are  %. 

"We  collected  a  large  series  of  adult  and  newly  transformed  Rana  aurora 
aurora  in  April  1933,  ]ust  north  of  Klamath,  Del  Norte  County.  It  was  raining 
and  frogs  were  abundant  on  the  sopping  forest  floor.  On  subsequent  examina- 
tion one  of  the  supposed  red-legged  frogs  turned  out  to  be  a  female  Ascaphtis" 
(W.  F.  Wood,  Calif.,  1939,  p.  no). 

Wallace  Wood's  individuals  that  transformed  in  April  interest  us.  We  early 
recorded  transformation  at  17-21  mm.  and  observed:  All  this  material 
(USNM  no.  337)  except  two  specimens,  26  and  44  mm.,  we  interpret  as  re- 
cently transformed.  The  National  Museum  frogs  were  taken  at  the  Columbia 
River  on  a  United  States  exploring  expedition,  and  Dr.  Barnhart  states  that 
the  expedition  came  to  the  Columbia  River  "late  in  April,  1841,"  and  arrived 
two  weeks  later  at  Puget  Sound.  After  our  varied  acquaintance  with  this  form 
we  conclude  that  these  and  Wood's  transformecs  must  have  come  from  early 
breeders. 

"Little  is  known  of  the  life-history.  The  eggs  are  laid  in  masses  during 
March  and  April  in  backwaters  of  lakes  and  streams.  Tadpoles  collected  in  a 
lake  near  Victoria,  B.C.  on  May  cjth,  1941,  when  about  i  inch  in  length,  trans- 
formed in  July  of  the  same  year"  (G.  C.  Carl,  B.C.,  194?,  p.  48). 

"Eggs:  The  mosaic  appearing  masses  are  made  up  of  large  eggs  with  a  vol- 
ume of  about  1.2  cc.  each.  The  jelly  is  very  transparent,  loose,  and  viscid.  Three 
envelopes  are  present  in  addition  to  the  vitclhnc  membrane,  which  is  easily 
discernible  in  most  cases.  Although  rather  indistinct  all  envelopes  can  be  seen 
without  the  aid  of  a  lens  when  viewed  in  proper  light.  Most  difficult  to  make 
out  is  the  middle  envelope,  which  is  the  thinnest.  The  fairly  large  vitellus  is 
black  at  the  animal  pole  and  creamy  white  at  the  vegetal.  Measurements  are: 
vitellus  3.04  mm.  (range  2.^1  to  3.56  mm.);  vitellinc  capsule  3.28  mm.  (range 
2.56  to  3.87  mm.) ;  inner  envelope  570  mm.  (range  4.0  to  6.68  mm.) ;  middle 
envelope  6.80  mm.  (range  6.25  to  7.93  mm.) ;  outer  envelope  1 1.</>  mm.  (range 
10.0  to  14.0  mm.). 

".  .  .  Several  differences  are  noticeable  between  the  eggs  of  /?.  a.  aurora  and 
its  very  near  relation  R.  a.  draytonii.  In  general,  eggs  of  R.  a.  aurora  are  larger; 
the  first  thing  to  strike  the  eye,  the  vitellus,  is  perceptibly  greater  as  is  the 


414  HANDBOOK  OF  FROGS  AND  TOADS 

dimension  of  the  outer  envelope.  The  outer  envelope  is  also  less  distinct  in 
R.  a.  aurora.  Vitelline  membranes  are  more  easily  seen  in  R.  a.  draytomt.  With 
R.  a.  draytonii  the  three  gelatinous  envelopes  become  progressively  greater  in 
thickness  from  the  inner  to  the  outer,  in  R.  a.  aurora  the  middle  envelope  is  the 
thinnest.  The  entire  egg  mass  of  R.  a.  aurora  is  from  two  to  four  inches  greater 
in  diameter  than  that  of  R.  a.  draytonii. 

"R.  a.  aurora  eggs  somewhat  resemble  those  of  R.  sylvattca.  The  looseness 
of  the  jelly  and  form  of  the  mass  are  similar,  but  the  lack  of  a  middle  envelope 
and  usually  smaller  dimensions  of  individual  eggs  and  masses  readily  separates 
R.  sylvattca  from  R.  a.  aurora"  (R.  L.  Livezey  and  A.  H.  Wright,  Gen.,  1945, 
pp.  702-703). 

Journal  notes:  March  27,  1942.  Stopped  between  Hoh  and  Borachiel  rivers, 
Olympic  Peninsula,  Wash.,  beside  a  wonderland  meadow,  with  spots  of  snow, 
Ledum,  cranberry,  etc.  Here  in  ditch  saw  several  fresh  R.  a.  aurora  egg  masses. 
Saw  one  composite  mass  of  frogs'  eggs  getting  stranded,  2  ft.  by  3  ft.  in 
diameter.  (Must  have  been  15  to  30  masses  here.  Mating  congress  in  one  place 
as  in  R.  sylvatica.)  In  each  egg  envelope  were  green  algae  as  in  salamander 
eggs- 

March  19,  Oregon.  From  11:00  A.M.  to  12:50  stopped  between  Hauser  and 
Lakeside.  In  a  series  of  pools  beside  a  road  in  a  wood  clearing  recently  cut  off 
were  several  pussy-willow  ponds.  One  has  cattails;  each  has  some  sedge  tus- 
socks. All  have  some  log  entanglements.  On  these  logs,  along  the  banks,  and 
in  shallow  mounds  in  the  pond  these  frogs  sometimes  sit,  not  infrequently 
two  in  a  place,  either  young  or  females.  Lots  of  vegetation  in  bottom  of  ponds. 
Centers  sometimes  3  feet  deep.  No  eggs.  All  adults  caught  are  females.  Where 
are  the  males?  The  females  all  ripe.  Some  of  the  young  are  bright  red  under 
legs  and  even  on  sides  of  pectoral  region.  Throats  much  spotted.  .  .  .  Later 
took  in  a  ditch,  i  foot  wide  but  2  feet  deep,  one  female  at  least  20  feet  from 
pond,  on  way  to  pond.  These  frogs  surely  look  like  wood  frogs,  and  are  wood 
frogs. 

March  20.  Up  McKenzie  R.  with  Ruth  Hopson.  .  .  .  Caught  little  R. 
aurora  and  saw  two  adults.  What  is  mass  of  Ambystoma-hkc  eggs  left  high 
and  dry? 

March  27.  Between  Nolan  Cr.  and  Hoh  River  found  beside  road  a  ditch  at 
least  4  feet  across,  containing  plenty  of  vegetation  and  with  sedge  clumps  on 
its  wooded  side.  Beside  sedge  clumps  are  Arnbystoma-like  masses.  In  shallow 
end  of  ditch  Anna  espied  frogs'  eggs,  quite  dark  in  color.  The  eggs  are  far 
apart  m  mass,  at  least  %  inch  from  each  other.  The  fresh  mass  is  quite  bluish 
in  cast,  and  in  water  6  inches  deep.  One  mass  at  least  8-10  inches  across!  An- 
other 6  inches  across!  Three  good  masses  and  a  partial  one;  must  be 
R.  a.  aurora.  Where  are  the  R.  a.  aurora  males  and  females?  Heard  seven. 
Stopped  beside  another  stream.  Here  found  two  Rana,  one  adult  with  much 
red  and  cloudiness  and  one  small  one,  yellow  below. 


FROGS:  RANIDAE 


415 


Plate  LXXXVII.  Rana  aurora  aurora,  i.  Plate  LXXXVIII.  Rana  aurora  draytonii 

Male  (X%)«  2.  Female  (X%).  3.  Female          (X1/^).  1,2,4.  Females.  3.  Male. 


416  HANDBOOK  OF  FROGS  AND  TOADS 

March  30.  Went  to  Sparaway  Lake  with  Mr.  James  Slater  and  later  to 
another  more  brushy  lake.  Here  he  gets  R.  a.  aurora.  Found  one  mass  of 
JR.  a.  aurora  eggs.  The  egg  has  two  egg  envelopes  with  a  very  visible  vitelline 
membrane.  The  tadpoles  have  three  rows  of  teeth  on  the  upper  labium  and 
four  rows  on  the  lower. 

Authorities  corner: 
H.  S.  Fitch,  Ore.,  1936,  p.  640. 
I.  McT.  Cowan,  B.C.,  1939,  p.  48. 
W.  C.  Brown  and  J.  R.  Slater,  Wash.,  1939,  pp.  17-18. 

"Some  previous  collectors  have  placed  specimens  of  this  species  [Rana  cas- 
cadae\  with  Rana  aurora  aurora.  .  .  .  Rana  aurora  aurora  and  Rana  cascadae 
overlap  a  little  in  distribution  in  the  Canadian  Zone"  (J.  R.  Slater,  Wash., 
I939b,p.  149). 

California  Red-legged  Frog,  Drayton's  Frog,  Bullfrog,  Long-footed  Frog, 

Bloody  Nouns,  Rocky  Mountain  Frog,  Western  Wood  Frog, 

French  Frog,  Leconte's  Frog 

Rana  aurora  draytonii  (Baird  and  Girard).  Plate  LXXXVIII;  Map  27. 

Range:  Coastal  California  to  Lower  California.  Few  records  in  Sierras  or 
east  of  them.  Introduced  in  Nevada  (Lmsdale).  "Inhabits  chiefly  the  Upper 
Sonoran  life-zone,  but  extends  into  Transition  and  Lower  Sonoran"  (J.  Grin- 
nell  and  C.  L.  Camp,  Calif.,  1917,  p.  149). 

Habitat:  Ponds,  lakes,  or  marshes.  Large  permanent  pools  or  water  courses. 
Linsdale  in  Lower  California  (L.  C.,  1932,  p.  353)  gives  several  habitats.  "More 
than  half  the  specimens  in  the  scries  listed  above  were  captured  in  or  at  the 
margins  of  streams.  One  individual  was  taken  from  a  spring,  one  from  a 
slough,  one  from  the  ground  beneath  willows  near  a  creek,  two  in  rat  traps 
at  the  edge  of  a  swamp,  three  from  a  marsh,  and  four  from  a  lake.  One  of  the 
frogs  was  captured  by  the  use  of  a  trout  fly." 

"It  is  known  to  occur  in  a  number  of  water-storage  reservoirs  and  other 
artificial  ponds,  and  it  has  been  cultivated  with  some  degree  of  success  in  ponds 
of  quiet  water  on  one  or  more  'frog  farms'  within  the  State.  ...  In  general, 
the  ponds  and  creek  pools  which  are  the  haunts  of  this  species  are  seldom 
frozen,  and  if  they  do  freeze,  the  ice  seldom  lasts  for  more  than  part  of ^ one 
day"  (T.  I.  Storer,  Calif.,  1925,  pp.  236-237). 

"This  is  the  large  frog  which  frequents  the  ponds  and  streams  on  both  sides 
of  the  mountains.  It  is  usually  found  where  water  is  permanent  and  when 
disturbed  takes  refuge  in  the  weeds  at  the  bottom.  It  has  been  collected  in 
Imperial  County.  It  may  be  distinguished  from  the  other  frogs  of  the  genus 
Rana  in  this  territory  (except  the  Leopard  frog)  by  the  ridges  on  the  sides  of 
the  body,  and  from  that  frog  by  the  absence  of  the  conspicuous  oval  blotches 
which  characterize  the  latter"  (L.  M.  Klauber,  Calif.,  i934a,  p.  7). 


FROGS:  RANIDAE  417 

Size:  Adults,  2^-5%  inches.  Males,  63-100  mm.  Females,  58-136  mm. 

General  appearance:  This  is  a  large,  stout,  rough-skinned  frog  with  a  thick 
broad  head.  It  is  the  largest  of  California's  native  frogs.  The  leg  bars  on  tibia 
and  femur  may  be  so  prominent  as  to  appear  zebrahke.  The  back  is  olive  drab 
or  buffy  brown,  spotted  with  light-centered  spots;  the  groin,  heavily  spotted; 
the  lateral  fold,  light  pinkish  cinnamon;  the  line  on  the  upper  jaw,  cream 
buff  or  pale  pinkish.  The  underside  of  the  hind  legs  and  the  inner  half  of 
the  tarsus  and  foot  are  pinkish  to  red.  The  young  are  apt  to  be  conspicuously 
spotted. 

Color:  Sentenac  Canyon,  San  Felipe  Cr.,  Calif.,  from  L.  M.  Klauber,  March 
25,  1928.  Female.  Upper  parts  Isabella  color,  Saccardo's  olive,  drab,  or  buffy 
brown.  Lateral  told  light  pinkish  cinnamon.  Line  on  upper  jaw  pale  pinkish 
cinnamon  or  cream-buff.  Brighter  than  in  male.  Around  tubercles  between 
lateral  folds  and  on  sides  are  black  rings  with  light  pinkish  cinnamon  in 
center  on  tubercle.  On  sides  this  becomes  congo  pink  or  dragon's  blood  red. 
Groin  is  dark  purplish  black  and  spotted  with  olive-buff  or  cartridge  buff. 
Femur  marked  with  dark  purplish  black  and  cartridge  buff  on  dorsal  sur- 
face. Underside  of  hind  legs  and  inner  half  of  tarsus  and  foot,  pale  pinkish 
cinnamon,  congo  pink  to  dragon's  blood  red.  Iris  orange. 

Male.  Upper  parts  citrine-drab  or  yellowish  olive  to  light  grayish  olive  on 
forearms;  more  uniform  on  back  with  few  black  spots  or  specks.  Groin  more 
spotted  than  in  female,  more  yellow  present.  Ventral  parts  more  heavily 
spotted  and  reticulated;  otherwise  as  in  female. 

Structure:  Thumb  enlarged,  slightly  two-lobed;  thumb  base  quite  large  in 
female  and  in  some  individuals  would  be  mistaken  for  a  male;  prominent 
ridge  from  posterior  angle  of  eye  to  shoulder;  skin  thick  and  roughened  with 
many  small  papillae,  even  the  eardrum  may  be  so  roughened;  eardrum  small; 
dorsolateral  folds  prominent;  sacral  hump  prominent;  back  with  regularly 
placed  light-centered  spots;  male  without  visible  vocal  sac. 

"This  species  resembles  very  much  ...  \R.  aurora\  in  its  external  appear- 
ance. It  differs,  however,  in  having  a  truncated  snout,  the  nostrils  consequently 
nearer  to  its  tip  than  to  the  eyes.  The  eyes  themselves  and  tympanum  are  pro- 
portionally larger  than  in  R.  aurora,  the  limbs  more  developed  and  the  tongue 
much  narrower.  The  ground  color  is  olivaceous  green,  maculated  with  black 
on  the  upper  region  of  the  body  and  limbs,  whilst  underneath  the  hue  is  uni- 
color,  except  sometimes  under  the  head,  breast  and  hind  legs,  where  the  brown 
and  white  mingle  in  circular  dots.  Specimens  were  collected  at  San  Francisco, 
California,  and  on  Columbia  River  by  Mr.  Drayton  himself,  to  whom  we 
take  pleasure  in  dedicating  this  species"  (S.  F.  Baird  and  C.  Girard,  Ore., 
1854,  p.  174). 

Voice:  "A  series  of  low  tremulous  or  'gurgling1  notes,  resembling  somewhat 
the  notes  uttered  by  Rana  boyhi"  (T.  I.  Storer,  Calif.,  1925,  p.  238). 

Breeding:  This  species  breeds  from  January  to  March.  The  eggs  are  laid 


418  HANDBOOK  OF  FROGS  AND  TOADS 

in  overflow  areas  of  permanent  pools,  the  mass  attached  to  vegetation.  The 
jelly  is  soft  and  viscid;  the  outline  of  individual  eggs  is  evident  on  the  sur- 
face. The  egg  is  Yi2  inch  (1.8-2.1  mm.),  the  envelopes  %,  %o,  %  inch  (3.5 
mm.,  4.4  mm.,  8.5  mm.),  the  eggs  black  and  white.  The  dark  brownish, 
mottled  tadpole  is  3%  inches  (83  mm.)  and  has  tooth  ridges  %.  The  larval 
period  is  5-7  months.  (Data  from  Storer,  1925.)  The  tadpoles  transform  from 
May  to  August  at  %-i%  inches  (18-30  mm.). 

This  frog  is  one  of  earliest  to  spawn  in  California,  the  crest  coming  in  Jan- 
uary and  February.  The  customary  size  at  transformation  is  probably 
22-30  mm.  Of  seven  specimens  taken  by  us  Aug.  14, 1917,  at  Oceanside,  Calif., 
none  was  smaller  than  32  mm.  and  all  were  past  transformation.  In  one  set, 
that  of  the  University  of  Michigan  (no.  65203),  appear  nine  transforming  frogs 
25-35  mm.  in  body  length,  with  tail  stubs  from  13-48  mm.  Two  specimens 
just  transformed  are  31  and  32  mm.  and  14  specimens  are  29-37  mm. 

In  egg  measurements  one  is  not  sure  whether  or  not  some  other  author 
interpreted  the  vitelline  membrane  as  an  envelope.  We  measured  some  of 
Storer's  eggs  from  Thornhill,  Calif.,  to  be  able  to  compare  his  statements  with 
other  material.  We  found  eggs  of  i.8-2.i-mm.  vitelh,  a  vitelline  membrane, 
and  three  envelopes,  the  outer  of  which  was  loose  and  the  two  inner  much 
firmer.  We  therefore  agree  with  Storer's  diagrams  and  measurements. 

Journal  notes:  Feb.  14,  1942,  Hoover  Ranch.  Cook  caught  me  an  adult  fe- 
male R.  a.  draytonii. 

March  10,  Stanford.  Anita  Daugherty  yesterday  gave  us  R.  a.  draytonii  eggs 
laid  in  lab  from  frogs  taken  at  Los  Trancos  Creek  March  i. 

April  12.  Out  with  Tracy  and  Ruth  Storer.  .  .  .  After  going  along  stream 
tributary  to  N.  Fork  Cosumnes  River  north  of  Plymouth,  Calif.,  near  the 
Amador-El  Dorado  County  line,  in  the  depressions  of  gold  workings,  saw  a 
R.  a.  draytonii  jump  into  a  deep  pool.  It  rested  on  the  bottom.  Brought  net 
in  front  of  it  and  poked  it  from  rear  as  I  pushed  forward  with  net.  Caught 
this  one  and  three  more.  One  was  in  grass;  it  leaped  into  a  clump.  Put  net 
between  clump  and  water.  Then  kicked  from  rear  and  in  jumped  my  frog. 
These  ugly  deposits  are  from  dredging  or  placer  workings. 

April  22.  With  Leo  T.  Hadsall  and  party  stopped  beside  drainage  ditch  on 
level,  near  Fresno,  Calif.;  some  three  or  four  R.  a.  draytonii  jumped  in. 

May  5,  La  Posta.  At  a  creek  beside  a  tule  area  saw  some  amphibian  jump 
in.  Had  to  know  whether  it  was  Bttfo  calif  ornictts,  my  search,  or  Rana  a. 
draytomi.  It  proved  the  latter.  Three  and  one-half  to  4  miles  south  of  Buck- 
man's  Spring  at  bridge  below  confluence  of  La  Posta  and  Cottonwood  Cr. 
took  one  R.  a.  draytontt.  Saw  no  Rana  tadpoles. 

Authorities9  corner: 
M.  C.  Dickerson,  Gen.,  1906,  p.  216. 
T.  I.  Storer,  Calif.,  1925,  p.  245. 
J.  M.  Linsdale,  Nev.,  1938,  p.  25. 


FROGS:  RANIDAE  419 

California  Yellow-legged  Frog,  Thick-skinned  Frog,  Boyle's  Frog 
Rana  boylii  boylii  (Baird).  Plate  LXXXIX;  Map  28. 

Range:  Western  Oregon,  northern  and  central  portions  of  California, 
chiefly  west  of  the  high  Sierra  Nevada. 

"Life  zones  Upper  Sonoran  and  Transition"  (J,  Grinnell  and  C.  L.  Camp, 
Calif.,  1917,  p.  146). 

"Except  for  a  small  area  in  southwestern  Oregon,  the  range  of  Rana  boylii 
boylii  jeems  to  lie  entirely  within  the  state  of  California.  It  includes  the  north- 
western part  of  the  state,  east  to  the  McCloud  River,  Shasta  County,  and  to 
the  western  foothills  of  the  Sierra  Nevada  below  4100  feet  altitude.  The  form 
occurs  also  at  Mono  Lake;  in  the  Sierras  it  has  been  taken  south  to  the  vicinity 
of  Walker  Pass,  Kern  County;  but  where  it  meets  the  range  of  R.  b.  muscosa 
along  the  coast  is  not  known.  Specimens  from  the  vicinity  of  Walker  Pass 
show  a  more  contrasted  pattern  of  coloration,  possibly  indicating  approach 
to  the  southern  subspecies,  described  below"  (C.  L.  Camp,  Calif.,  1917,  p.  118). 

Habitat:  Margins  of  springs,  streams,  and  fresh  water  lakes  (J.  Grinnell  and 
C.  L.  Camp,  Calif.,  1917);  gravelly  streams  (K.  Gordon,  Ore.,  1939). 

"It  is  confined  to  the  immediate  vicinity  of  permanent  streams,  at  least  those 
where  water  holes  persist  through  the  dry  season.  It  is  most  common  along 
streams  having  rocky  beds,  but  occurs  also  in  ones  having  mud  buttoms"  (H. 
S.  Fitch,  Ore.,  1936,  p.  640). 

"This  little  frog  inhabits  the  slow  flowing  streams  of  the  coastal  areas,  and 
may  be  found  in  considerable  numbers  in  the  semipermanent  pools,  formed 
as  the  streams  become  low  at  the  end  of  the  rainy  season"  (J.  R.  Slevin,  B.C., 
1928,  p.  139). 

At  Stevens  Creek  near  Palo  Alto,  Calif.,  in  shallow  and  quiet  areas  we 
found  small  1?.  boylii.  Up  one  or  two  little  runs  bordered  by  Rhits  (poison 
oak),  saw  four  more. 

T.  I.  Storer  says:  "The  adult  frogs  of  this  species  spend  much  of  their  time 
perched  on  rocks  in  the  stream  or  on  the  bank,  but  in  the  latter  place  they 
never  go  more  than  two  or  three  feet  from  the  margin  of  the  water.  If  ap- 
proached, even  from  the  direction  of  the  stream,  they  invariably  seek  safety 
by  leaping  into  the  water,  and  immediately  swim  with  swift  strokes  down 
to  the  bottom.  In  streams  with  silt  on  the  bottom  they  hide  in  the  mud  and 
silt  which  their  movements  stir  up;  in  clear  waters  they  take  refuge  under 
overhanging  rocks." 

Size:  Adults,  i%~3  inches.  Males,  39-67  mm.  Females,  40-75  mm.  It  is  the 
smallest  of  the  three  subspecies  of  Rana  boylii. 

General  appearance:  This  is  a  small  frog  with  stout,  broad  body.  The  skin 
on  the  back,  legs,  and  tympanum  is  thick  and  rough  with  small  brownish 
papillae.  The  color  of  the  back  is  black,  light  gray,  greenish,  or  brownish  with 
indistinct  dark  mottling.  There  is  a  patch  of  lighter  color  on  top  of  the  head 


420 


HANDBOOK  OF  FROGS  AND  TOADS 


with  a  darker  area  behind  crossing  the  upper  eyelids.  The  tympanic  region 
is  darker  than  the  head.  Red  is  never  present  in  the  coloration.  The  venter  is 
white  with  pale  yellow  on  the  posterior  part  and  on  the  hind  legs.  The  throat 
and  sides  of  the  body  are  mottled  with  black. 

Of  four  adult  specimens  of  R.  b. 
boylii  taken  April  2,  1942,  only  one,  a 
male,  had  a  tendency  to  be  spotted  in 
the  groin.  All  had  spotted  throats. 
Three  males  had  short,  light  tubercular 
ridges  back  of  the  angle  of  the  mouth. 
It  was  either  a  continuous  fold  or  two 
to  four  tubercles  in  a  row;  then  came  a 
hiatus  and  then  another  ridge  or  patch 
of  one  to  three  tubercles.  The  skin  was 
rougher  than  in  Rana  pretiosa  or  R.  a. 
aurora.  Quite  often  they  had  a  red  line 
along  the  dorsolateral  fold,  the  lower 
belly  lumiere  green,  underside  of  femur 
mustard  yellow,  then  primuline  yellow, 
then  yellow  ocher,  becoming  in  rear 
portion  aniline  yellow. 

Color:  Female.  Mill  Valley,  Muir 
Woods,  Calif.,  from  John  Needham, 
March  18, 1930.  Dorsal  color  in  general 
is  buffy  olive,  light  brownish  olive, 
citrine-drab,  or  olive  lake,  coming  to  be, 
in  the  intervals  between  the  leg  and 
arm  crossbars,  Isabella  color,  clay  color, 
or  honey  yellow  on  the  tarsus  and  outer 
edge  of  the  foot.  The  crossbars  are  Sac- 
cardo's  olive.  On  rear  of  femur  and  in 
the  groin  are  several  rather  prominent 
black,  olive,  or  deep  olive  areas.  On  the 
rear  of  femur  the  dark  areas  have  ecru- 
olive  or  light  yellowish  olive  intervals.  A  ridge  from  rear  of  upper  labial  margin 
around  angle  of  mouth,  a  short  area  back  of  eye  and  irregularly  along  back 
where  dorsolateral  fold  would  be,  and  patches  on  either  side  of  knee  have 
orange-cinnamon  tubercles.  The  front  of  the  tibia  bears  an  empire  yellow  line 
separating  dorsal  from  ventral  surface.  The  top  of  the  head  from  front  of  one 
eyelid  to  the  other  and  forward  on  snout  is  Isabella  color  or  old  gold.  Ventral 
surfaces:  The  central  rear  half  of  the  femur  is  papillate,  orange,  or  deep  chrome, 
bounded  behind  by  a  more  or  less  prominent  black  or  deep  olive  band.  In  front 
of  this  orange  area  and  on  lower  belly  is  a  wash  of  picric  yellow.  Some  of  same 


Map  28 


FROGS:  RANIDAE 


421 


color  on  groin,  on  rear  of  tibia,  on  venter  of  arm,  and  in  axil  of  arm.  The  top  of 
the  foot  has  a  wash  of  empire  yellow.  The  throat  and  breast  are  white  to  pale 
pinkish  buff  with  orange-cinnamon  spots  on  throat  and  vaguely  outlined  on 
breast.  These  become  verona  brown  spots  on  edge  of  lower  jaw.  Along  side 
as  back  merges  into  belly  are  picric  yellow  spots  among  the  deep  olive.  The 
eye  is  black,  almost  obscured  by  numerous  specks  of  buff-pink  or  vinaceous- 
pink  on  lower  portion  giving  place  to  light  green-yellow  and  pale  orange- 
yellow  specks  on  the  upper  half.  Pupil  rim  in  lower  half  partakes  of  color 
of  specks  of  that  part,  and  so  also  the  rim  on  the  upper  part. 

Structure:  Head  bioad  and  pointed;  web  of  foot  large,  only  slightly  scal- 
loped; toes  blunt,  tips  slightly  expanded;  thumb  of  male  enlarged,  the  swelling 
having  two  lobes;  tympanum  small,  distinct. 

"Rana  boyht,  Baird. — A  broad  depressed  ridge  of  skin  on  each  side  of  back. 
Skin  finely  tubercular  above.  Head  broader  than  long.  Tympanum  scarcely 
evident,  pustulated.  Tibia  more  than  half  the  length  of  body;  hind  foot  less 
than  half  this  length;  webbed  entirely  to  the  horny  tips;  outer  toe  decidedly 
longer  than  the  third.  An  elongated  tubercle  at  base  of  inner  toe,  with  another 
opposite  to  it.  Above  dull  reddish  olivaceous,  with  indistinct  blotches  on  the 
back,  and  fascia  on  the  legs.  Beneath  yellowish,  mottled  anteriorly.  Two  inches 
long.  Hab.  California  (interior)"  (S.  F.  Baird,  Gen.,  1856,  p.  62). 

"Those  characters  which  pertain  to  this  subspecies  alone  are:  hind  leg  long, 
inside  angle  of  bent  tarsus  reaching  at  least  to  narcs  and  usually  beyond  when 
leg  is  advanced  along  body;  tibia  elongate,  reaching  usually  beyond  anus  when 
flexed  and  held  at  right  angles  to  axis  of  body;  fourth  toe  on  reflexed  hind  foot 
never  reaching  beyond  end  of  knee  and  often  not  quite  to  fold  of  skin  below 
knee;  head  broad  and  pointed  when  viewed  from  above,  its  width  two  and 
one-third  to  two  and  two-thirds  times  in  body-length;  skin  on  back,  legs  and 
tympanum,  thick  and  rough  with  minute  brownish  spines;  color  dorsally 
varying  from  nearly  uniform  to  black  to  light  gray,  greenish  or  brownish,  with 
darker  markings,  if  present,  usually  indistinct;  there  is  always  a  patch  of 
lighter  color  on  top  of  head  between  nares  and  eyes,  and  behind  this  a 
darker  area  crossing  posterior  half  of  each  eyelid  and  merging  insensibly  be- 
hind into  the  general  dorsal  coloration"  (C.  L.  Camp,  Calif.,  1917,  pp.  117- 
118). 

"Size  small;  head  broader  than  long,  depressed;  snout  rounded,  projecting 
beyond  mouth;  canthus  rostralis  distinct;  loreal  region  concave;  nostrils  nearer 
to  tip  of  snout  than  to  orbit;  distance  between  nostrils  equals  interorbital  width. 
Interorbital  width  less  than  width  of  upper  eyelid.  Tympanum  small,  distinct, 
covered  with  small  tubercles,  about  one-half  diameter  of  eye.  Fore  limbs  mod- 
erately robust;  digits  rather  long,  first  and  second  equal,  third  much  the  long- 
est; no  lateral  fold  along  sides  of  fingers;  subarticular  tubercles  small  or 
moderately  large,  rounded,  prominent;  outer  metatarsal  tubercles  elongate, 
prominent,  inner  metatarsal  tubercle  rounded,  somewhat  obscure;  web  full, 


422  HANDBOOK  OF  FROGS  AND  TOADS 

extending  to  tip  of  longest  toe.  Skin  rugose,  covered  with  warts  or  tubercles 
on  back  and  sides;  posterior  surface  of  the  thigh  covered  with  small  tubercles; 
dorso-lateral  fold  obscure;  sacral  hump  rather  prominent;  a  well  developed 
fold  from  lip  to  side  of  neck  or  shoulder.  Vomerine  teeth  in  two  oblique  series, 
widely  separated  anteriorly,  between  and  a  little  behind  the  choanae"  (J.  R. 
Slevin,  B.C.,  1928,  p.  137). 

Voice:  No  description  of  the  call  is  recorded.  The  frog  has  internal  vocal 
sacs. 

Breeding:  This  frog  breeds  from  the  latter  part  of  March  to  the  first  of  May. 
The  egg  mass,  in  shallow  water  toward  the  margin  of  streams,  is  attached  to 
sides  of  stones  in  the  stream  bed  and  is  like  a  compact  cluster  of  grapes,  the 
individual  eggs  on  the  surface  distinct,  the  jelly  firm.  The  eggs  are  more 
closely  set  than  are  those  in  a  R.  ptpiens  mass. 

"Three  envelopes  present.  Outer  envelope  3.88-4.47  mm.,  ave.  4.0  mm,;  jelly 
firm.  Middle  envelope  2.58-3.35  mm.,  ave.  2.8  mm.  Inner  envelope  2.32-2.94 
mm.,  ave.  2.5  mm.  All  envelopes  distinct.  Vitellus  1.93-2.48  mm.,  average  2.2 
mm.;  black  above  and  white  below.  Mass  2  by  2  by  i%  to  2  by  4  by  2%  inches 
in  dimensions.  Eggs  firmly  attached  to  each  other.  909-1037  eggs  in  a  mass. 
Deposited  in  shallow  water  near  the  margins  of  streams,  attached  to  stones 
in  stream  bed.  Pick  up  much  sediment"  (R.  L.  Livezey  and  A.  H.  Wright, 
Gen.,  1947). 

The  tadpole  is  medium,  2  inches  (50  mm.),  deep  olive  in  color,  and  with 
tooth  ridges  %,%,%•  After  3  to  4  months,  the  tadpoles  transform  from  June 
to  September,  at  %-iYs  inches  (20-30  mm.). 

"Evans  Creek  (Oregon)  on  April  29,  1934.  •  •  •  Numerous  egg  masses  at- 
tached to  large  pebbles  and  newly  hatched  tadpoles  were  seen.  The  spawning 
places  were  in  pools  four  or  five  inches  deep  where  slow  currents  or  eddies 
caused  constant  circulation  of  water  around  the  egg  masses"  (H.  S.  Fitch, 
Ore.,  1936,  p.  640). 

A  frog  taken  by  J.  C.  Bradley,  June  1907,  Blue  Lake,  Humboldt  Co.,  Calif., 
was  29  mm.  and  past  transformation;  and  in  a  scries  of  six  from  Marin  Co. 
by  J.  C.  Bradley,  which  were  22.5-30  mm.,  none  had  arm  scars.  Transforma- 
tion of  some  of  this  group  must  have  come  at  20  mm.  or  smaller.  H.  C.  Kellers, 
April  13,  1913,  secured  two  specimens  (USNM  nos.  49874-5)  25  and  31  mm. 
These  doubtless  wintered  over  from  the  previous  year.  This  same  collector 
has  taken  this  species  in  Calistoga,  Napa  Co.,  Calif.,  as  small  as  23  mm.  Mr. 
Rutter,  Oct.  17,  1899,  in  Mariposa,  Calif.,  secured  two  specimens  (nos. 
38830-31)  29.0  and  25.5  mm.  These  might  be  of  the  season.  The  specimens 
(USNM  nos.  20895-20903)  secured  by  H.  W.  Henshaw  Oct.  17, 1893,  at  Ukiah, 
Mendocino  Co.,  Calif.,  ranging  from  20-29  mm->  maY  als°  be  of  the  same 
season. 

April  8,  1942.  Ate  at  Trail,  Ore.  Went  up  the  creek.  In  a  spring  area  saw 
several  R.  boylii  boylii.  Took  in  side  pools  small  R.  b.  boylii.  They  swim  out 


FROGS:  RANIDAE  423 

into  deep  water  and  then  generally  back  to  edge  or  to  shallow  water.  In  the 
middle  of  stream  riffles  saw  an  adult  R.  b.  boylii  swimming.  While  waiting 
for  Bert  to  search  above  the  bridge  for  more  snakes  (Thamnophis  o.  hydro- 
phila)  I  went  to  stream  side  near  bridge.  Across  the  stream  I  saw  what  I 
thought  an  adult  R.  b.  boylii.  This  was  a  crossing  place  in  the  stream  with  flat 
rocks;  just  above  were  rapids  and  just  below,  deep  pools  with  rapid  current 
in  water  falling  into  pools.  I  waded  across  stream  catching  sight  of  two  adult 
frogs.  Then  in  a  shallow  side  pocket  with  water  flowing  through  but  not 
rapidly,  attached  to  a  broken  notch,  was  a  mass  of  R.  b.  boyln  eggs.  The  water 
was  3  to  4  inches  deep.  The  egg  cluster  was  somewhat  plinthlike.  Eggs  were 
fresh  and  looked  black.  Bert  came  down  and  reported  that  this  must  be  type 
locality  for  Fitch's  T.  o.  hydrophila.  No  end  of  adult  Rana  b.  boylii  leaped  into 
midstream  ahead  of  him.  It  is  interesting  to  see  them  helplessly  carried  over 
the  shallow  rock  bottom  by  the  current  until  they  find  a  deep  groove.  They 
are  shy  frogs  and  you  can't  get  within  catching  distance  before  they  dash  for 
the  bottom  under  the  edge  of  a  near-by  stone  if  there  is  a  handy  one. 

Journal  notes:  Feb.  14,  1942.  Trip  to  Theodore  Hoover  ranch,  Calif.  Went 
up  steep  side  of  small  creek  and  in  it  Rodgers  secured  a  yellow-legged  frog. 
Around  sawmill  in  little  shallow  pools  found  several  small  R.  boylii  boylii. 
During  the  day  the  boys  got  me  one  or  two  more. 

March  15.  Trip  north.  In  Tuttle  Ranch,  Calif.,  took  R.  b.  boylii. 

March  18.  Started  at  10:30  along  Smith  River  road  toward  Smith  River 
village,  Calif.  It  is  the  east  segment  of  the  triangle.  Along  a  bank  covered  with 
ferns  and  oxalis  in  a  rill  where  water  goes  under  a  road,  Anna  caught  a  male 
R.  b.  boylii.  In  pool  at  other  end  of  pipe  I  took  two  more. 

March  23.  With  Kenneth  Gordon  and  group  to  Santiam  River,  Ore.  In 
ponds  beside  stream  under  rock,  Storm  and  Livezcy  took  three  R.  boylii  boylti. 
Under  board,  Anna  found  a  female. 

April  12.  Out  with  Tracy  and  Ruth  Storer.  Region  of  dredging  and  placer 
working,  north  of  Plymouth,  Calif.  Over  in  the  stream  Tracy  and  Ruth  Storer 
and  Anna  kept  seeing  yellow-legged  frogs  leap  into  the  water.  This  is  a  creek 
with  plenty  of  nlHy  spots.  In  a  fast-water  stretch  Tracy  Storer  found  a  mass 
of  R.  b.  boylii  eggs  attached  to  stone  3  or  4  feet  from  bank  in  water  10  inches 
deep,  a  plinth.  Before  we  got  through  we  found  five  more  masses— one  very 
fresh  and  hardly  expanded,  this  less  plinthlike. 

April  1 8.  Went  to  Hasting's  Natural  History  Reservation  to  sec  Jean  Lins- 
dale.  He  has  H.  regtlla  and  R.  boylii  there. 

Authorities9  corner: 
C.  E.  and  M.  D.  Burt,  Ariz.,  1929,  p.      H.  S.  Fitch,  Ore.,  1936,  pp.  640-641. 

432.  H.  S.  Fitch,  Ore.,  1938,  p.  148. 

J.  Grinnell,  J.  J.  Dixson,  J.  M.  Linsdale, 

Calif.,  1930,  p.  143. 

"There  is  thus  a  good  chance  that  on  the  desert  slope  of  the  San  Gabriel 


414  HANDBOOK  OF  FROGS  AND  TOADS 

Mountains  the  frogs  are  boylii  rather  than  muscosa'  (C.  L,  Camp,  Calif.,  1917, 
p.  120).  [A  statement  verified  by  Marr  26  years  later.] 

"In  a  herpetological  collection  made  along  the  San  Gabriel  River,  San 
Gabriel  Mountains,  Los  Angeles  County,  August  n,  1940,  I  have  four  speci- 
mens of  Rana  boylii  boylii  and  none  of  Rana  boylii  muscosa.  ...  It  is  barely 
possible  that  this  frog  has  been  introduced  by  some  chance  into  the  San  Gabriel 
River"  (}.  C.  Marr,  Calif.,  1943,  p.  56). 

Sierra  Madre  Yellow-legged  Frog,  Southern  Yellow-legged  Frog 

Rana  boylii  muscosa  Camp.  Plate  XC;  Map  28. 

Range:  San  Gabriel,  San  Bernardino,  and  San  Jacinto  Mountains  of  south- 
ern California.  "This  form  inhabits  stream  canons  in  southern  California 
south  of  Tehachapi  Pass.  Localities  of  record  extend  from  Pacoima  Canon 
near  San  Fernando  eastward  and  southward  along  the  southern  and  western 
slopes  of  the  San  Gabriel,  San  Bernardino  and  San  Jacinto  ranges  as  far  as 
Keen's  Camp,  in  the  latter  range.  It  has  been  found  also  at  Little  Rock  Creek 
on  the  north  slope  of  the  San  Gabriel  Mountains.  The  range  in  altitude  is  from 
1200  feet  near  Sierra  Madrc  to  6500  feet  on  Fish  Creek  in  the  San  Bernardino 
range"  (C.  L.  Camp,  Calif.,  1917^  p.  119). 

"Southern  California  south  o£  Ventura  County"  (J.  R.  Slevin,  Calif.,  1934, 

p.  4»). 

"The  previously  recorded  southern  limit  of  this  species  has  been  Keen 
Camp  in  the  San  Jacinto  Mountains,  Riverside  County,  California.  Recently 
it  has  been  found  to  be  plentiful  in  Pauma  Creek,  where  it  flows  through 
Upper  Doane  Valley,  Palomar  Mountain,  San  Diego  County,  at  elevation  5100 
feet.  The  first  specimens  at  this  location  were  taken  by  Dr.  E.  H.  Taylor  of 
the  University  of  Kansas  during  a  collecting  trip  in  which  the  writer  partici- 
pated" (L.  M.  Klauber,  Calif.,  1929,  pp.  15-16). 

Habitat:  "Occupies  the  upper  Sonoran  and  Transition  life  /ones.  Lives 
along  streams  in  narrow  rockwalled  canons"  (J.  Grinnell  and  C.  L.  Camp, 
Calif.,  1917,  p.  148).  "More  or  less  common  in  streams  throughout  (Los 
Angeles)  County"  (C.  M.  Bogert,  Calif.,  1930,  p.  5). 

Size:  Adults,  i  %~3/4  inches.  Males,  44-66  mm.  Females,  47-85  mm.  Our 
material  taken  in  Santa  Anita  Canyon  and  Snow  Creek  in  spring,  1942,  falls 
into  three  classes:  young  33-40  mm.,  males  49-56  mm.,  and  females  49- 
64  mm. 

General  appearance:  "Like  Rana  boylii  boylti,  but  attaining  much  larger 
size,  and  (except  in  young)  with  no  light  patch  in  front  of  dark  areas  across 
upper  eyelids.  Dorsal  ground  color  usually  lighter  than  in  R.  b.  boylti,  light 
yellow  to  brown,  contrasting  with  the  darker  mosslike  patches  on  the  back. 
Tips  of  toes  more  expanded  than  in  boylii9  (C.  L.  Camp,  Calif.,  1917,  p.  119). 

Color:  Male.  San  Gabriel  Canyon,  Los  Angeles  region,  Calif.,  from  Berry 


FROGS:  RANIDAE 


4*5 


Plate  LXXXIX.  Rana  boylii  boylii.  i.  Plate  XC.  Rana  boylii  muscosa 

Male  (X%).  2,3.  Females  (X%).  1,2-  Females.  3,4.  Males. 


426  HANDBOOK  OF  FROGS  AND  TOADS 

Campbell,  April  n,  1930.  Upper  parts  from  tip  of  snout  back  between  eyes 
and  lateral  folds  to  vent  ecru-olive  or  deep  olive-buff.  Costal  folds  dark  olive- 
buff.  Interspaces  on  hind  legs  dark  olive-buff  instead  of  ecru-olive.  Color  below 
costal  fold  olive-buff  or  ivory  yellow  and  same  on  under  parts  and  front  half 
of  under  part  of  hind  legs.  An  oblique  spot  from  eye  to  angle  of  mouth,  one 
from  tympanum  obliquely  backwards,  colonial  buff.  On  front  of  femur,  top 
of  foot,  axilla,  slightly  on  head  and  front  edge  of  arm  deep  colonial  buff. 
Throat  cartridge  buff  with  fine  buffy  olive  spottings.  Spot  on  top  of  head  and 
back  brownish  olive.  Same  color  also  on  hind  legs  and  forelegs,  on  sides  be- 
coming buffy  olive.  Underside  of  head  and  hind  ventral  half  of  femur  pale 
grayish  vinaceous  or  pale  vinaceous-fawn.  Thumb  excrescences  clove  brown, 
also  underside  of  hand. 

Female.  Upper  Doane  Valley,  Palomar  Mt.,  San  Diego  Co.,  Calif.,  from 
L.  M.  Klauber  and  Dr.  E.  H.  Taylor,  Sept.  2,  1928.  Upper  parts  from  tip  of 
snout  back  between  eyes  and  two  costal  folds  to  vent  grayish  olive  or  citrine- 
drab  (lighter  than  in  male).  Costal  fold  dark  to  deep  olive-buff,  color  below 
costal  fold  pale  olive-buff,  and  the  same  on  under  parts.  An  oblique  spot  from 
eye  to  angle  of  mouth,  one  back  of  angle,  one  over  arm  insertion  and  one  in 
axilla  primrose  yellow  or  olive-buff.  Under  parts  of  legs  like  venter.  Where 
lighter  color  joins  upper  color  considerable  of  deep  colonial  buff  or  chamois. 
This  color  comes  strongly  into  light  intervals  between  crossbars  on  hind  legs 
and  somewhat  on  forelegs.  Some  wash  of  same  color  on  throat,  which  is  not 
as  spotted  as  in  male.  Sides  below  costal  folds  heavily  spotted  with  brownish 
olive.  Spots  on  dorsum  from  tip  of  snout  to  vent  smaller  and  often  more  or 
less  olive-ocher  encircled.  Dark  line  from  eye  to  nostril  to  labial  margin.  Black 
spot  or  bar  below  eye.  Edge  of  lower  jaw  black  or  brownish  olive  and  colonial 
buff  or  pale  olive-buff.  Top  of  fingers  deep  colonial  buff  or  chamois.  Dark 
bars  of  hind  legs  brownish  olive  or  olive  with  some  grayish  olive  spots  in  the 
dark  color.  Hind  webs  dark  with  some  chamois.  Horizontal  bar  of  black  or 
olive  through  eye;  vertical  bar  of  same  below  pupil;  interspaces  between  the 
horizontal  and  vertical,  isabella  color  or  even  primrose  yellow;  upper  portion 
of  eye  colonial  buff  with  black  dot  at  the  top.  Sometimes  at  forward  edge  of 
eye  occurs  pale  yellow-green. 

Structure:  Tips  of  toes  more  expanded  than  in  R.  b.  boylii;  dorsolateral  fold 
indistinct,  not  pitted  anteriorly;  tympanum  and  area  surrounding  it  very 
rough,  covered  with  small  tubercles;  web  of  hind  foot  extending  nearly  to  tips 
of  toes,  broader  in  expanse  than  in  R.  b.  boyhi;  swelling  on  thumb  of  male  has 
two  lobes,  the  constriction  being  diagonal  not  transverse. 

"Vomerine  teeth  on  two  oblique  ridges  between  nares;  head  pointed  in  out- 
line as  viewed  from  above,  broad,  its  width  entering  body-length  two  and 
two-thirds  times;  hind  limbs  long,  posterior  side  of  bent  tarsus  reaching  for- 
ward to  snout;  fourth  toe  on  hind  foot  reaching  forward  not  quite  to  knee- 
fold;  dorsolateral  fold  indistinct,  not  pitted  anteriorly;  tympanum  and  area 


FROGS:  RANIDAE  427 

surrounding  it  very  rough,  beset  with  small  tubercles,  web  of  hind  foot  ex- 
tending nearly  to  tips  of  toes;  outer  and  inner  metatarsal  tubercles  distinct; 
plantar  tubercles  very  large;  tips  of  toes  expanded,  disc-shaped;  distal  end  of 
flexed  tibia  held  at  right  angles  to  body  reaching  anus"  (C.  L.  Camp,  Calif., 
1917,  p.  119). 

"Ground  color  above  light  gray  or  yellowish,  with  numerous  black  blotches 
or  reticulation  on  head,  body  and  limbs.  Posterior  surface  of  thigh  rich  yellow. 
Under  surfaces  yellowish  or  whitish,  gular  region  slightly  spotted  or  marbled 
with  gray"  (J.  R.  Slevin,  Gen.,  1928,  p.  141). 

Voice:  Nothing  on  record. 

Breeding:  This  species  breeds  during  April.  The  eggs  are  not  yet  described. 
The  tadpole  is  much  like  that  of  R.  boylii  sierrae  and  reaches  61  mm.  in  length. 
Its  body  is  rather  flat,  the  tail  musculature  heavy  and  almost  uniform  in  width 
for  about  half  the  length  of  tail.  Tooth  ridges  are  variable,  for  example,  %  in 
one,  %  in  another,  /4  in  another.  Possibly  they  winter  as  tadpoles  and  trans- 
form in  the  spring.  They  transform  at  about  %  inch  (20-24  mm.). 

Two  tadpoles,  collected  Sept.  5,  1922,  in  San  Jacinto  Mountains,  Dark 
Valley,  by  Prof.  J.  C.  Bradley  are  41  and  43  mm.,  not  fully  grown.  The  crests 
remain  about  the  same  to  the  point  where  musculature  tapers. 

One  specimen  from  near  Clarcmont,  Calif. ,  1907,  is  30  mm.,  just  beyond 
transformation.  The  two  smallest  in  the  National  Museum  (nos.  54899- 
54900),  taken  by  E.  J.  Brown  in  San  Gabriel  Canyon,  Los  Angeles  Co.,  Calif., 
in  February,  1916,  are  24  and  26  mm.  Apparently  they  were  transformed  the 
season  before.  This  seems  to  imply  transformation  in  the  neighborhood  of 
20  mm. 

On  June  20,  1928,  Storer  secured  at  BlufI  Lake,  3400  feet  elevation,  San 
Bernardino  Mts.  and  Co.,  a  tadpole,  61  mm.  long,  body  23  mm.  Its  tail  was 
rounded;  apparently  the  tadpole  was  approaching  transformation;  its  tooth 
formula  was  %•  Other  specimens  had  tails  that  were  not  broad,  crests  quite 
spotted.  Some  were  practically  transformed  at  23  and  24  mm. 

On  May  i,  1909,  C.  L.  Camp  found  specimens  20-28  mm.;  on  Feb.  17,  1918, 
J.  Shaw  took  a  25-mm.  specimen  and  on  May  30,  1918,  a  29-0101.;  on  Nov.  13, 
1937,  W.  F.  Wood  took  specimens  from  24-36  mm.  One  wonders  if  they  winter 
over  as  tadpoles. 

Journal  notes:  April  26,  1942,  Calif.  Started  for  Santa  Anita  Canyon  about 
9:30  A.M.,  hazy  over  the  mountains.  Rode  about  3  miles  up  canyon  to  public 
parking  spot.  Went  down  trail  to  the  river,  about  %  mile  down,  and  then 
a  mile  or  so  up  the  river.  Near  where  trail  went  to  ranger  cabin  crossed  a 
very  rocky,  steep  gorge  with  small  amount  of  swift  water.  It  was  dark  and 
shady  at  this  point.  We  saw  no  frogs  there.  As  we  reached  the  river  another 
side  stream  came  in,  White  Creek.  We  went  slowly  up  the  river.  At  one  spot 
beside  the  river  but  not  connected  with  it  was  a  pool,  a  rather  scummy,  dark- 
looking  pool,  small,  perhaps  4  by  6  feet.  Here  Bert  caught  one  male  and  one 


428  HANDBOOK  OF  FROGS  AND  TOADS 

small  Rana  b.  muscosa  and  some  newts,  Triturus  torosus,  and  found  quantities 
of  beautiful  newt  egg  clusters  attached  to  sticks.  More  were  under  a  board  and 
some  were  attached  to  a  board  lying  at  the  end  of  the  pool.  (Caught  two 
H.  aremcolor.}  Much  granitic  rock  here — light  gray  in  color — almost  no  sedi- 
ment in  the  stream.  On  our  return,  Bert  stopped  at  White  Creek  and  there 
caught  three  female  R.  b.  muscosa ,  nice  large  ones.  He  carried  them  in  a  large 
wet  bag.  He  had  two  newts  with  them.  When  we  reached  the  car  he  put  the 
bag  into  the  thermos  bottle  with  a  little  water.  Either  too  much  water  or  too 
much  newt  killed  two  of  the  females  by  the  next  day.  A  male  JR.  b.  muscosa 
leaped  into  a  quiet  algae-covered  pool  filled  with  brush.  Took  young 
R.  b.  muscosa  in  this  pool.  In  a  quiet  cutofl  saw  a  female  leap  into  creek  and 
go  under  a  stone.  I  was  too  leisurely  about  it.  Lost  it.  I  was  angry  to  lose  the 
female;  saw  no  more  except  two  or  three  young. 

On  return  came  to  Winter  Creek.  Here  on  a  sandy  bar  was  one  female.  It 
leaped  into  a  quiet  pool  near  the  base  of  a  falls.  Another  halfway  up  on  falls 
leaped  in  and  lodged  under  a  stone  part  way  down.  Caught  it.  In  a  pool  at 
top  of  this  sloping  granitic  falls  took  another  female  in  brushy  edge.  Many 
newts  mating  in  this  creek.  A  few  trout  in  Winter  Creek. 

April  30.  Trip  to  Idyllwyld.  At  mouth  of  Strawberry  Creek  in  cattail  area 
in  one  quiet  pool  saw  two  Rana  b.  muscosa.  Took  one  in  tules  below  bridge, 
and  later  above  bridge  there  leaped  in  from  a  boulder  a  female  H.  aremcolor. 
Went  to  Idyllwyld — no  luck.  Tried  several  places. 

May  9,  Palm  Springs.  Went  up  Snow  Creek  5  miles.  Here  road  ends  where 
water  company's  possessions  are.  The  Sierra  Club  of  Los  Angeles  was  meeting 
here.  Some  eight  to  ten  people  were  along  the  creek  awaiting  their  leader's 
coming.  A  beautiful  cool  mountain  stream.  There  used  to  be  a  fish  hatchery 
in  the  stream.  At  once  back  of  car — 1850  feet  elevation — we  saw  R.  boylit 
muscosa;  the  first  near  weeds  at  edge  leaped  among  some  rocks  but  came  up, 
as  they  frequently  do,  and  was  easy  of  capture. 

May  9,  Snow  Creek.  These  frogs  were  most  frequently  seated  on  the  rocks. 
I  would  flop  the  net  flat  on  the  rock  and  the  frog  would  leap  into  the  netting. 
Sometimes  they  disappeared  before  I  reached  close  enough  to  catch  them. 
Occasionally  I  could  approach  close  and  seize  them  with  my  left  hand.  We  saw 
only  two  or  three  small  ones.  I  collected  about  five  females  before  I  got  a  male. 
In  the  end  there  were  seven  females  and  two  males  in  the  lot.  A  fine  grayish- 
white,  bouldery,  tumbling  stream  of  cold  water.  No  wonder  these  muscosa 
are  much  lighter  than  the  sterrae  and  boyhi.  They  are  a  shade-loving  form  and 
have  a  decided  odor.  They  sat  on  damp  rocks  above  the  water  line.  The  first 
male  was  darker  than  the  rest. 

Authorities9  corner: 
C.  L.  Camp,  Calif.,  1917,  pp.  119-120. 
L.  M.  Klauber,  Calif.,  19343,  p.  7. 


FROGS:  RANIDAE  429 

Sierra  Nevada  Yellow-legged  Frog,  Western  Frog,  Pacific  Frog,  Stink  Frog 

Rana  boylii  sierrae  Camp.  Plate  XCI;  Map  28. 

Range:  California.  Southern  half  of  Sierra  Nevada,  above  7000  feet  altitude 
from  Yosemite  National  Park  on  the  north  to  southern  Tulare  County  on 
the  south. 

Habitat:  Meadows,  streams,  and  lakes  from  7000  to  10,500  feet  in  Yosemite 
Park,  and  to  11,500  feet  near  Mount  Whitney. 

"In  some  of  the  lakes  it  was  found  in  great  numbers,  appearing  as  soon 
as  the  ice  had  melted  in  late  June.  The  tadpoles  were  at  this  time  of  large  size 
and  must  have  been  hatched  from  eggs  ot  a  laying  not  more  recent  than  the 
previous  year"  (C.  L.  Camp,  Calif.,  1917!},  p.  122). 

Size:  Adults,  i%~3%  inches.  Males,  44-72  mm.  Females,  48-85  mm.  The 
measurements  of  19  males  and  30  females  in  the  California  Academy,  Stanford 
University,  and  University  of  California  collections  gave  a  range  of  44-65  mm. 
for  males  and  52-85  mm.  for  females;  average  male  55.5,  mean  56  mm.;  aver- 
age female  60  mm.,  mean  59,  66,  70  mm. 

General  appearance:  This  frog  is  like  Rana  boyln  boyln,  but  lacks  the  light 
patch  on  the  head.  The  back  is  slightly  roughened,  is  dark  olive  in  color  with 
some  darker  spots  showing  when  it  is  wet.  The  under  parts  arc  white  on  the 
throat,  creamy  or  buff  on  the  belly  and  the  underside  of  the  legs.  The  throat 
is  marked  with  black  spots.  The  sides  are  light,  heavily  mottled  with  distinct 
black  blotches.  The  jaw  is  heavily  mottled  with  light  and  dark.  There  is  a 
distinct  dark  blotch  on  the  upper  part  of  the  forearm.  The  inner  surface  of 
the  arms  is  light,  outer  surface  greenish,  marked  with  dark  blotches.  The  legs 
are  marked  with  dark  spots.  The  angle  of  the  jaw  forms  a  swollen  light  area 
marked  with  one  or  two  dark  spots,  and  back  of  this  is  another  light  tubercle 
or  swollen  area.  The  head  is  short  and  broad  with  the  nostrils  far  apart,  snout 
short,  eyes  small  and  widely  separated.  Costal  folds  low  and  inconspicuous. 
The  young  look  something  like  newly  transformed  green  frogs. 

"With  the  general  characters  of  Rana  boyln  boylii,  but  hind  leg  usually 
shorter  and  head  relatively  narrower;  tympanum  smoother;  and  light  patch 
on  top  of  head  wanting"  (C.  L.  Camp,  Calif.,  igi7b,  p.  120). 

Color:  Female.  Yosemite  Park,  from  C.  A.  Harwell,  Oct.  12,  1929.  (The 
frog  was  cold  when  described,  hence  dark.)  Back  deep  olive,  citrine-drab, 
light  brownish  olive,  few  black  or  dark  spots  centered  with  old  gold  between 
lateral  folds  from  arm  insertion  level  to  vent.  More  heavily  spotted  on  sides 
below  lateral  folds  and  spots  sometimes  light-centered  there  where  belly  color 
and  spots  are  reticulate.  Something  of  same  spotting  on  front  of  dorsum  of 
femur  and  rear  of  forelimb.  Throat  and  upper  and  lower  labial  edge  heavily 
spotted  with  black.  A  very  light  white  or  cartridge  buff  line  begins  below  eye 
and  extends  across  angle  of  mouth  or  just  above  it  to  arm  insertion.  Some- 
times this  is  interrupted  below  tympanum,  then  resumed  diagonally  down 


430  HANDBOOK  OF  FROGS  AND  TOADS 

from  tympanum,  then  interrupted  to  appear  as  a  swollen  patch  above  arm 
insertion.  The  posttympanal  and  arm  insertion  cartridge  buff  patches  may 
have  two  or  three  fine  black  spots  on  them  and  considerable  black  extending 
upward  between  them  from  arm  insertion.  Upper  jaw  and  face  pale  olive- 
buff  with  numerous  black  spots.  Tympanum  clear,  like  background  of  dorsum 
or  top  of  head.  Background  on  hind  legs  and  forelegs  dark  olive-buff  or  old 
gold;  mottled  or  spotted  with  black  or  olive;  lateral  fold  of  same  color.  Under- 
side of  hind  legs  and  forelegs  pale  chalcedony  yellow,  cream  color,  or  seafoam 
yellow;  a  tinge  of  same  color  on  lower  belly.  On  tarsus  old  gold  or  aniline 
yellow.  Throat  and  pectoral  region  white.  Underside  of  forefeet  and  hind  feet 
light  brownish  drab  or  hair  brown;  iris  black  dotted  with  citron  to  honey 
yellow,  or  strontium  yellow;  pupil  rim  broken  in  front  and  behind. 

Male.  Lake  Alpine,  7300  ft.,  Alpine  Co.,  Calif.,  from  Tracy  I.  Storer,  July 
29, 1930.  Back  buffy  olive,  citrine-drab,  or  grayish  olive.  Under  the  lens  it  may 
look  deep  olive-buff.  On  sides  interspace  color  is  deep  olive-buff,  ivory  yellow, 
or  cartridge  buff,  becoming  chamois  on  edges  of  belly.  The  interspaces  of 
hind  legs  and  of  forelegs  are  light  brownish  olive  or  almost  Isabella  color. 
Dorsolateral  fold  in  front  half  isabella  color  or  cinnamon-brown  to  tawny 
or  sayal  brown.  Large  spots  of  bister  or  auburn.  Whole  of  interspaces  and 
spots  covered  by  spaced  black  points.  On  sides,  dorsolateral  fold,  and  top  of 
femur  these  surmount  larger  papillae.  Forearm,  front  of  femur,  rear  of  tibia, 
tarsus,  and  foot  with  transverse  bars  of  same  color  as  large  dorsal  spots.  Lower 
rear  half  of  femur  with  many  papillae  tipped  with  reed  yellow  or  deep  colonial 
buff.  Throat  to  line  of  arm  insertion  and  upper  and  lower  labial  fold  heavily 
spotted  with  deep  olive  or  citrine-drab.  Middle  belly  reed  yellow,  its  sides 
chamois.  Underside  of  hind  legs  and  forelegs  colonial  buff  or  cream-buff.  In- 
side of  tarsus  and  lower  end  of  tibia  chamois.  Rear  half  of  venter  of  femur 
vinaceous-drab  or  deep  brownish  vinaceous,  each  papilla  tipped  with  deep 
olive-buff  or,  better,  reed  yellow  or  deep  colonial  buff.  Underside  of  fingers 
and  of  toes  like  rear  half  of  venter  of  femur.  Cream  buff  or  colonial  buff  fold 
from  below  eye  to  angle  of  mouth  broken  below  tympanum,  broken  again, 
resumed  as  spot  above  arm  insertion.  Iris  rim  picric  yellow  or  lemon  yellow 
above.  Iris  mars  orange  or  English  red. 

Structure:  Hind  legs  shorter;  head  relatively  narrower;  tympanum 
smoother  than  in  Rana  b.  boyhi;  web  of  hind  foot  very  large,  extending  to 
tips  of  toes;  tips  of  toes  not  much  expanded;  dorsolateral  fold  strongly  pitted 
anteriorly;  swelling  on  thumb  of  male  bilobate. 

"Vomerine  teeth  rudimentary,  on  two  oblique  ridges  nearly  meeting  be- 
tween and  slightly  behind  nares  .  .  .  ;  head  viewed  from  above  rounded  in 
outline;  head-width  contained  three  times  in  body-length;  hind  limbs  short, 
posterior  side  of  bent  tarsus  reaching  forward  to  anterior  corner  of  eye;  fourth 
toe  on  hind  foot  reaching  forward  to  end  of  bent  knee;  dorso-Iateral  fold 
indistinct,  strongly  pitted  anteriorly;  tympanum  nearly  smooth,  with  scat- 


FROGS:  RANIDAE  431 

tered  hispid  points;  web  of  hind  foot  very  large,  extending  to  tips  of  toes; 
outer  metatarsal  tubercle  rudimentary,  inner  one  small;  plantar  tubercles 
small;  tips  of  toes  not  much  expanded;  distal  end  of  flexed  tibia,  held  at  right 
angles  to  body,  just  reaching  anus;  color  above,  dark  yellowish  brown,  ob- 
scurely marked  with  indefinite  darker  vermiculations;  lower  surface  yellow, 
faintly  dotted  with  brown  beneath  chin;  upper  lip  below  eye  mottled;  no 
dark  cheek  patch;  hind  limbs  not  distinctly  barred  with  dark  bands. 

"This  seems  to  be  the  subspecies  of  boyln  which  approaches  most  closely 
to  the  species  pretiosa"  (C.  L.  Camp,  Calit.,  1917^  pp.  120-122). 

Voice:  No  data.  No  vocal  sacs.  Croaks  not  very  loud  in  captive  males. 

Breeding:  As  soon  as  ice  has  melted  in  the  mountain  lakes,  these  frogs  breed 
in  June  and  July.  The  eggs  are  approximately  like  those  of  Rana  b.  boylii  (T.  I. 
Storer,  Calif.,  1925).  The  tadpole  is  fairly  large,  2%  inches  (72  mm.),  its  body 
flattened,  the  tail  musculature  wide  for  an  inch  or  more,  then,  suddenly  ta- 
pering, the  tail  tip  rounded  01  spatulate.  The  crests  are  broader  toward  the  tip 
than  at  the  body.  The  tooth  ridges  are  %,  %,  or  %.  The  period  of  develop- 
ment is  i  year,  the  animal  passing  the  winter  as  a  tadpole.  It  transforms,  dur- 
ing July  and  August,  %-i  %  2  inches  (20-27  mm.) . 

In  the  summer  of  1907,  Prof.  J.  C.  Bradley  at  Giant  Forest,  Calif.,  took 
transforming  young  measuring  21-26  mm.  On  Aug.  22-27,  1917,  we  took  at 
Alta  Meadows,  Sequoia  National  Park,  a  series  of  transforming  frogs  which 
measured  from  21  mm.  to  27  mm.,  the  average  and  mode  being  23  mm.  Of 
our  twelve  females,  three  had  ripe  eggs  to  be  laid. 

At  Davis,  Calif.,  we  examined  some  of  Professor  Storer's  material.  He  had 
males  54-72  mm.  and  females  58-82  mm.  in  length.  He  had  a  young  specimen 
36  mm.  long  July  14,  and  others,  near  transformation,  20,  21.5,  22,  22  mm. 
July  3,  1930.  One  tadpole  of  54  mm.  (body  length  21.5  mm.)  had  a  tail  that 
tended  to  be  rounded,  even  spatulate,  at  the  tip.  Its  teeth  were  %.  Those  of 
another  were  %.  The  eggs  had  three  jelly  envelopes;  the  vitellus  was  2.0  mm.; 
and  the  egg  was  about  7  mm.  across.  Egg  masses  were  attached  to  the  banks  of 
a  small  creek. 

"One  mass  of  eggs  was  collected  by  Osgorxl  R.  Smith  on  April  23,  1942  at 
Convict  Creek,  Mono  County,  California,  at  an  elevation  of  7,000  feet.  This 
clutch  was  attached  to  stems  of  sedge  and  contained  about  120  eggs.  .  .  . 

"Mass  plinth-like,  about  28  X  40  mm.,  individual  eggs  one-quarter  to  one- 
half  inch  apart.  There  are  besides  the  vitellinc  capsule  three  gelatinous  en- 
velopes, all  clear  and  transparent.  The  outline  of  the  egg  is  distinct  and  the  firm 
jelly  does  not  collect  much  debris.  All  envelopes  arc  quite  distinguishable  in 
good  light  without  a  lens,  as  is  the  vitelline  capsule.  Least  in  thickness  is  the 
middle  envelope,  the  next  in  thickness  is  the  inner,  being  about  two  times  the 
thickness  of  the  middle,  and  the  thickest  is  the  outer  which  is  approximately 
twice  that  of  the  inner.  The  vitellus  is  black  at  the  animal  pole  and  grey-tan 
at  the  vegetal.  Measurements  are:  vitellus  2.17  mm.  (range  1.81  to  2.30  mm.); 


432  HANDBOOK  OF  FROGS  AND  TOADS 

vitclline  capsule  2.58  mm.  (range  2.25  to  2.75  mm.) ;  inner  envelope  3.93  mm. 
(range  3.75  to  4.81  mm.) ;  middle  envelope  4.60  mm.  (range  4.25  to  5.00  mm.) ; 
outer  envelope  7.19  mm.  (range  6.43  to  7.87  mm.). 

"Measurements  on  the  eggs  preserved  before  they  had  time  to  attain  their 
full  size  showed  the  vitellus  to  average  somewhat  less  (av.  1.88  mm.)  than 
the  other  eggs  taken  from  natural  surroundings"  (R.  L.  Livezey  and  A.  H. 
Wright,  Gen.,  1945,  p.  703). 

Journal  notes:  Aug.  23, 1917,  Alta  Meadows,  Calif.  In  the  pools  on  the  mats 
were  plenty  of  Rana  boylii  sierrae.  They  stink  like  minks  or  other  weasels. 
They  are  very  slow.  One  can  run  a  net  under  them  or  put  a  net  on  the  bottom 
in  front  and  touch  the  back  parts.  They  are  very  yellow  underneath.  The 
species  is  almost  through  transformation.  The  larvae  look  on  the  venter  some- 
what like  large  toad  tadpoles.  We  found  a  few  transformed  individuals. 

On  June  5,  1930,  O.  E.  Van  Deman,  while  looking  for  Bufo  canonis  on 
Peregoy  Meadows,  Yosemite  Park,  found  several  frog  egg  masses  in  the 
stream  flowing  through  the  meadow.  From  his  description  (in  1931)  it  seems 
likely  that  they  were  this  form. 

July  23, 1934,  Aspen  Valley  and  White  Wolf,  Yosemite  Valley.  In  a  boulder- 
strewn  stream  found  Rana  boylii  sierrae.  In  a  pool  with  brush  saw  a  frog  put 
out  and  stay  poised.  You  are  much  impressed  with  the  spotted  condition  and 
the  stripe  on  upper  jaw.  The  type  of  habitat  is  not  unlike  that  of  Rana  tara- 
htimarac  except  that  this  is  high  wooded  country  and  a  running  mountain 
stream  whereas  Rana  tarahttmarae  is  in  desert  canyons.  They  surely  belong 
in  the  same  group. 

Middle  Fork  crossing  Tuolumne  River,  8000  feet.  In  clear  stream  on  riffles, 
in  edges  of  pools,  or  among  grassy  clumps  are  Sierra  yellow  legs.  They  sit 
rather  flattened  close  to  stone  or  bottom.  They  permit  close  approach  and  are 
easy  of  capture.  Some  are  quite  light  and  then  are  very  conspicuous,  but  if 
dark  they  may  be  hard  to  see  on  the  bottom.  Males  have  thumbs  with  two 
pads  and  throats  clear  or  spotted.  One  male  very  brilliant  orange  or  cadmium 
orange  on  belly  and  hind  legs,  becoming  xanthine  orange  on  rear  of  thighs 
and  on  the  toes.  In  bag  the  males  gave  a  low  croak.  Throat  pale  orange  yellow. 
Females  when  held  in  hand  may  give  a  clicking  or  chuckling  note.  Under 
parts  of  females  paler.  When  the  male  croaks,  just  the  center  of  the  throat 
swells  out  slightly.  Tioga  Pass — in  one  spot  50  tadpoles  in  one  scoop,  25  in  the 
next  try. 

April  22,  1942.  Went  up  King's  River  road  with  L.  T.  Hadsall  and  party. 
In  Sycamore  Cr.  in  quiet  portions  of  the  stream  we  saw  little  R.  b.  sierrae.  The 
adults  would  leap  from  banks  or  gravelly  strands  into  the  very  swift  water 
and  were  often  hard  to  catch,  particularly  if  it  was  a  bouldery  portion  of  the 
creek.  No  eggs  to  be  seen  in  the  stream.  We  caught  several  small  females  and 
two  males  unbelievably  small.  These  we  put  together  in  the  fish  pail.  The  very 


FROGS:  RANIDAE 


433 


Plate  XCI.  Rana  boylit  sierrae.  1,3.  Fe-  Plate  XCll.   Rana  capita.    1,5.   Males 

males  (X%).  2.  Male  (Xtf).  (X*)-  2-  Male  (X%>.  3-  Male  croaking 


434  HANDBOOK  OF  FROGS  AND  TOADS 

little  males  mated,  and  one  female  through  this  last  week  (April  22-29) 
laid  a  few  eggs. 

Authorities'  corner: 
C.  L.  Camp,  Calif.,  igiyb,  p.  123. 
J.  Grinncll  and  T.  I.  Storer,  Calif.,  1924,  p.  664. 

Florida  Gopher  Frog,  Gopher  Frog,  Snake  Frog,  Southern  Gopher  Frog, 

Florida  Frog 

Rana  capita  Le  Conte.  Plates  XCII;  XV;  Map  26. 

Range:  North  Carolina  to  Florida. 

"In  the  writer's  possession  is  an  adult  male,  collected  in  May  1919  by  Charles 
E.  Snyder  near  Pinelands,  Hampton  County,  South  Carolina,  and  it  is  quite 
likely  that  the  species  extends  a  good  deal  further  north  and  east  than  even 
this  locality'*  (R.  F.  Deckert,  Fla.,  1920,  p.  26).  [Note:  Pineland,  Jasper  Co., 
is  the  locality  in  question? — E.  B.  Chamberlain,  S.C.,  1939,  p.  28.] 

Habitat:  Almost  solely  in  the  burrows  of  the  gopher  turtle,  which  is  com- 
mon in  the  higher  pine  barrens  and  sandy  hills.  They  breed  in  cypress  and 
open  ponds.  We  saw  one  go  into  a  small  opening  which  looked  like  a  rat  hole. 
The  hole  extended  18  inches  into  the  ground  and  the  end  was  only  9-12  inches 
below  ground.  There  we  found  a  female  gopher  frog.  Near  another  turtle's 
burrow  we  saw  a  similar  hole  with  a  smooth  worn  spot  about  8-12  inches 
away,  the  resting  place  of  the  frog. 

The  residents  of  Chesser  Island,  Ga.,  assured  us  that  these  frogs  betake 
themselves  to  rathke  holes,  hollow  stumps,  and  holes  beneath  stumps,  trees, 
and  logs.  Where  gopher  turtles  arc  absent,  they  might  resort  to  crayfish  bur- 
rows, as  Gaige  tells  us  Rana  areolata,  a  closely  related  species,  does.  Doubt- 
less no  other  frog  proves  quite  so  typical  of  the  higher  pine  barrens  unless  it 
be  the  oak  toad.  Occasionally  the  toad,  the  spadefoot,  and  the  swamp  cricket 
frog  (Pseitdacris  nigrita)  may  be  found  in  the  same  plant  association. 

"The  specimen  mentioned  here  was  caught  in  a  cage  trap  set  among  the 
sand  hills  to  catch  small  rodents"  (R.  F.  Deckert,  Fla.,  1920,  p.  26). 

"Moderately  common  on  the  oak  ridges  remaining  in  or  near  the  burrows 
of  Gophems  polyphemus  (Daudin),  going  to  ponds  only  to  breed"  (O.  C. 
Van  Hyning,  Fla.,  1933,  p.  4). 

"High  pine,  turkey  oak  ridges,  and  rosemary  scrub.  .  .  .  Fairly  common, 
but  secretive  and  ungregarious;  sometimes  breeding  in  very  large  numbers 
in  isolated  ponds.  .  .  .  Nocturnal,  but  occasionally  seen  seated  at  the  mouths 
of  burrows  on  dark  days.  Although  they  occupy  the  holes  of  Gopherus  poly- 
phemus extensively,  they  are  by  no  means  confined  to  such  retreats.  Wright 
(1933)  mentions  finding  them  in  small  burrows  which  he  took  to  be  those 
of  a  rat.  J.  D.  Kilby  took  one  in  the  bottom  of  the  burrow  of  a  Peromyscus 
polionottts;  I  have  twice  found  them  in  the  burrows  of  the  same  mouse,  and 


FROGS:  RANIDAE 


435 


have  also  seen  them  frequently  in  stump  holes  and  in  the  bottom  of  post  holes, 
whence  escape  was  apparently  impossible.  I  caught  one  yearling  in  the  mouth 
of  a  crayfish  burrow  at  the  edge  of  a  cypress  pond.  They  range  some  distance 
from  their  retreats  in  foraging  at  night,  although  Wright  .  .  .  believes  they 
spend  much  time  resting  on  *a  little  clear  place  —  a  short  distance  from  the 
hole.'  I  once  saw  one  seated  on  a  fallen  pine  log;  looking  about  I  located  a 
gopher-hole  some  thirty  feet  from  the  frog;  when  I  advanced  and  kicked  one 
end  of  the  log  the  frog  hopped  about  wildly  for  a  moment,  then  headed  di- 
rectly for  the  gopher-hole  and  disappeared  down  it.  Since  there  was  no  other 
burrow  in  the  immediate  vicinity,  and  since  the  hole  was  not  visible  from 
the  frog's  original  position,  it  seems  to  me  justifiable  to  interpret  this  behavior 
as  the  result  of  homing  instinct.  Several  observers  have  noticed  the  tendency 
of  capita  to  crouch  at  the  approach  of  an  intruder.  I  found  one  which  had  been 
stepped  on  and  killed  by  dogs  running  a  fox"  (A.  F.  Carr,  Jr.,  Fla., 


Size:  Adults,  2%-4%  inches.  Males,  68-101  mm.  Females,  77-108  mm. 

General  appearance:  This  large,  heavy-bodied  gray  frog  is  broad  forward 
and  has  a  slender  waist.  It  has  a  cavernous  mouth  and  prominent  eyes.  The 
gray  of  the  back  is  finely  speckled  with  black  dots  and  marked  with  three  or 
four  rows  of  round  dark  spots  between  and  often  crossing  the  dorsolateral 
folds.  There  are  many  more  along  the  sides.  The  arms  and  legs  are  con- 
spicuously barred;  and  the  throat,  breast,  and  much  of  the  underside  of  the 
hind  legs  are  heavily  dotted  with  dark.  The  males  have  yellow  on  folds, 
tubercles,  iris,  axilla,  and  groin,  but  the  females  have  little  or  no  yellow. 

Color:  Male.  Okefinokee  Swamp,  Ga.,  July  20,  1921.  Sides  of  head,  dorsum 
of  limbs  and  sides  below  the  costal  fold  French  gray  or  lilac  gray.  Between 
the  costal  folds  about  four  rows  of  black  spots.  Fore  and  hind  legs  with 
numerous  black  bars,  broken  at  times  or  speckled  with  French  gray.  Costal 
fold  from  honey  yellow  to  mustard  or  buff  (yellow  to  greenish  yellow  depend- 
ing on  degree  of  dark  specking  on  it).  Another  similar  stripe  along  upper 
jaw  and  over  arm  insertion.  All  the  tubercles  on  the  back  with  same  color. 
Under  parts  white  with  hair  brown,  fuscous,  or  purplish;  black  spots  on  throat 
and  chin.  Posterior  faces  of  hind  limbs  purplish  vinaccous.  Upper  part  of  iris 
black  then  buff  yellow  or  honey  yellow  over  the  pupil,  black  in  front  and  be- 
hind the  pupil,  two  light  gray  areas  beneath  pupil  separated  by  black. 

Female.  Okefinokee  Swamp,  Ga.,  July  2,  1922.  General  color  on  sides  French 
gray,  cinereous,  or  pale  purplish  gray.  Lateral  fold,  canthus  rostralis,  upper 
eyelid,  beneath  eye,  and  fold  back  of  angle  of  mouth  cream-buff  or  tilleul 
buff.  Ground  color  of  back  a  combination  of  colors  of  sides  and  lateral  fold. 
On  the  back  are  four  more  or  less  irregular  rows  of  black  rounded  spots,  three 
such  rows  on  the  sides.  Black  crossbars  on  limbs.  Spots  on  the  throat  bister  to 
snuff  brown.  Those  on  sides  of  belly  army  brown.  Underside  of  feet  and  hands 
and  femur  deep  vinaceous-lavender.  Iris  black  with  prominent  bar  of  colonial 


436  HANDBOOK  OF  FROGS  AND  TOADS 

buff  or  tilleul  buff  above  the  pupil;  below  the  pupil  may  be  two  small  bars  of 
French  gray,  cinereous,  pale  purplish,  or  pallid  purplish  gray.  In  another  frog 
the  two  lower  bars  are  united.  Rest  of  eye  black  with  a  little  dotting  of  the  two 
colors. 

Structure:  Wedge-shaped  body,  wide  in  head  and  slender  in  waist;  tuber- 
cles on  back;  thumbs  of  males  somewhat  enlarged  and  darkened;  males  with 
vocal  sacs  from  angle  of  mouth  halfway  to  the  groin. 

"Above  very  rough,  dark  grey  or  slate-color  speckled  with  black  with  six 
rows  of  roundish  spots  on  the  back;  side  speckled  and  irregularly  marked  with 
spots  of  the  same  form  and  color;  from  the  orbits  to  beyond  the  middle  of  the 
body  runs  a  broad  raised  line  or  cutaneous  fold;  and  another  from  the  corner 
of  the  mouth  to  the  insertion  of  the  arm.  Beneath  smooth,  yellowish  white, 
speckled;  spotted  and  varied  with  dusky;  top  of  head  coarsely  punctured,  back 
and  sides  tuberculous.  Head  very  large,  broad  and  blunt,  a  deep  concavity 
between  the  nostrils  and  the  eyes.  Indes  golden  mixed  with  black.  Tympanum 
of  the  color  of  the  body.  Lower  jaw  with  a  small  protuberance  or  point  re- 
sembling a  tooth.  Arms  and  legs  above  grey,  speckled  and  barred  with  black, 
beneath  yellowish  spotted  and  varied  with  dusky,  the  yellowish  color  more 
decided  at  the  axillae  and  groins.  Hind  part  of  the  thighs  granulate.  Fingers 
slightly  palmate  at  the  base,  the  first  longer  than  the  second.  The  second  toe 
twice  as  long  as  the  first.  Length  4.2  in.,  width  of  the  head  at  the  corners  of 
the  mouth  1.5  in.,  arm  1.87,  leg  4.75,  thigh  i.i  in.,  tibia  1.45,  foot  2.2. 

"Inhabits  Georgia  in  the  ditches  of  the  rice-fields"  (Le  Conte,  Gen.,  1856, 
p.  425). 

"This  singular  form  may  be  known  at  once  by  the  short  and  squat  form  of 
the  body  as  compared  with  the  size  of  the  head,  resembling  in  this  some  of 
the  Australian  Cystignathidae.  .  .  .  The  only  specimen  of  this  subspecies 
which  I  have  seen  is  the  following:  Rana  areolata  aesoptis  Cope.  Catalogue 
number  474?,  No.  of  spec.  i.  Locality,  Micanopy,  Fla.  From  whom  received: 
Dr.  T.  H.  Bean.  Nature  of  specimen:  alcoholic"  (E.  D.  Cope,  Gen.,  1889,  pp. 
412-413). 

Voice:  The  call  is  a  deep  hollow  roll,  intermediate  between  a  snore  and  a 
groan,  yatvh  h  h  h  ht  yatvh  h  h  h  h. 

The  croaking  is  a  very  remarkable  performance.  A  lateral  pouch  is  inflated 
on  each  side  of  the  neck  and  keeps  swelling  posteriorly  until  it  extends  half- 
way down  the  side  of  the  body,  attaining  a  size  nearly  equal  to  that  of  the 
frog's  head.  The  creature's  form  is  then  strongly  suggestive  of  a  batfish.  The 
note  and  likewise  the  inflation  lasts  for  a  second  or  so.  At  the  start  the  intervals 
between  notes  may  range  from  10  seconds  to  nearly  half  a  minute,  but  as  the 
frog  warms  to  its  performance,  a  croak  may  be  given  every  2  or  3  seconds. 
The  sound  will  probably  carry  a  quarter  of  a  mile.  When  we  first  heard  the 
note,  one  of  the  residents  thought  it  a  woodpecker,  but  I  recorded  that  it 
seemed  more  like  Rana  sphenocephala  in  tone,  though  the  latter  is  a  more 


FROGS:  RANIDAE 


437 


rasping  croak.  The  call  of  Rana  sphenoccphala  is  like  a  rattle  compared  with 
the  uniform  continuous  roar  of  Rana  capita.  In  chorus  it  sounded  to  Mrs. 
Wright  and  myself  like  the  surf  or  a  deep  guttural  roar  (possibly  a  trill  at 
times)  like  rolling  r's  down  deep  in  the  throat.  The  chorus  seemed  to  go  in 
waves  with  decided  crests. 

"When  the  interior  of  the  large  glass  jar  in  which  it  lives  is  sprinkled  and 
after  this  certain  noises  are  made  like  rustling  papers  or  water  running  from 
a  tap  this  frog  'sings/  The  call  resembles  a  loud  snore  similar  to  that  of  Rana 
pipiens,  but  much  coarser  and  louder.  During  the  calls,  which  are  repeated 
about  every  two  seconds  and  are  of  from  three  to  five  seconds'  duration,  the 
vocal  vesicles  over  the  arms  are  distended  into  hemispheres  about  the  size  of 
large  hazel  nuts"  (R.  F.  Dcckert,  Fla.,  1920,  p.  26). 

Breeding:  This  species  breeds  throughout  the  year.  The  eggs  are  in  a  large 
plmthlike  mass  about  6  by  8  inches  and  i%  inches  deep.  The  egg  is  Vi2  inch 
(2  mm.),  the  outer  envelope  %  inch  (5.2  mm.),  the  inner  %  inch  (3.8  mm.). 
They  hatch  in  4-4%  days.  The  tadpole  is  large,  $%  inches  (84  mm.),  full  and 
deep  bodies,  the  venter  strongly  pigmented,  the  tail  long.  The  color  may  be 
greenish,  with  yellowish  on  the  top  and  side  of  the  head,  the  belly  yellowish. 
The  tooth  ridges  are  %.  After  85  to  100  days,  the  tadpoles  transform  from 
August  to  Nov.  i,  at  iHa-1%  inches  (27-58  mm.). 

On  Aug.  17,  1922,  we  found  several  egg  masses  laid  in  a  cypress  pond.  On 
the  north  edge  in  water  6-8  inches  deep  and  30  feet  from  the  edge  of  the  pond, 
I  found,  i  inch  below  the  surface  at  the  base  of  a  small  cypress  (10  feet  high) 
and  among  some  brush,  a  mass  attached  to  an  upright  twig.  The  black  and 
cream  eggs  look  larger  and  farther  apart  than  are  those  of  Rana  sphenocephda. 
Near  a  large  cypress  stump  in  an  open  grassy  area  where  pine  cones  were  on 
the  bottom,  a  large  mass  was  attached  to  a  sedge  stem.  Its  top  was  level  with 
the  surface  of  the  water.  The  water  was  9  inches  deep.  The  mass  was  4  by 
5  inches  and  1%  inches  thick.  At  first  the  mass  impressed  all  of  us  as  bluish, 
and  two  of  us  independently  likened  it  to  R.  sylvatica  in  this  respect.  The 
lower  pole  may  be  cream  at  first,  but  it  soon  becomes  white.  The  whole  mass 
when  turned  over  gives  the  same  white  mass  impression  that  R.  sphenocephala 
and  R.  pipiens  egg  masses  give.  Another  mass  was  6  by  8  by  1%  to  2  inches 
thick. 

"The  writer  has  also  collected  this  species  during  its  breeding  season  in 
February,  near  Jacksonville,  Fla.  in  1912"  (R.  F.  Dcckert,  Fla.,  1920,  p.  26). 

"March  13  to  November  3  in  grassy  ponds.  They  must  travel  some  distance 
to  congregate  at  widely  separated  ponds,  since  I  have  found  them  a  mile  from 
water"  (A.  F.  Carr,  Fla.,  1940^  p.  64). 

Journal  notes:  Okefinokee  Swamp,  Ga.  In  its  normal  all-the-year  habitat, 
the  sand  hills  and  turtle  burrows,  these  frogs  are  seldom  seen  unless  one  de- 
liberately seeks  them.  They  usually  rest  at  the  mouth  of  the  burrow,  some- 
times a  foot  or  so  down  the  decline;  more  rarely  they  may  be  a  foot  or  more 


438  HANDBOOK  OF  FROGS  AND  TOADS 

from  the  incline,  on  little,  clear,  smooth  places  6  inches  in  diameter  a  short 
distance  from  the  hole.  One  area  we  visited  frequently.  From  midforenoon 
to  midafternoon,  they  were  seldom  out.  For  example,  on  July  2,  1922,  we 
visited  this  area  at  noon  and  2:30  P.M.  and  found  only  one  out,  but  at  mid- 
night we  readily  found  eight.  My  journal  reads,  Called  again  at  12  midnight. 
Francis*  special  friend  not  to  be  seen.  Anna's  friend  was  under  a  log,  the  en- 
trance a  foot  or  more  away  because  we  had  dug  it  back  from  under  the  log. 
Suddenly  something  tumbled  from  under  the  log  into  the  hole  before  I 
realized  what  had  happened.  Francis  found  another  at  a  hole  near  by.  I  found 
one  and  Miles  saw  three:  one  was  1-1%  feet  from  the  entrance.  This  one  I 
caught.  It  is  a  female.  Tried  for  one  or  two  more  but  did  not  succeed.  They 
are  not  so  strongly  held  by  light  as  are  most  Anura. 

July  n,  1922.  At  7:15  P.M.  we  started  for  the  gopher-frog  area  near  Chesser 
School.  Found  seven  or  eight  at  entrances  to  holes  or  a  foot  or  two  away.  At 
one  hole  Miles  saw  a  cotton  mouse  (Peromyscus  gossypinus) .  At  another  I 
saw  a  cotton  rat  (Sigmodon).  We  captured  three  frogs.  I  caught  one  by  hand. 
Could  have  taken  three  or  four  more.  Miles  took  the  net  without  its  handle, 
interposed  it  between  frog  and  hole.  Frog  like  a  woodchuck  makes  for  the 
hole  and  jumps  into  the  net.  Sometimes  this  works,  sometimes  it  does  not.  At 
first  I  espied  a  frog  at  the  entrance  and  tried  to  steal  up  on  it  from  opposite 
the  incline.  It  heard  me.  Wonder  if  one  could  catch  them  with  baited  hook 
and  line.  One  was  obliging  and  allowed  us  to  take  its  picture.  First  one  this 
year.  It  turned  around  twice  during  the  process,  then  popped  in.  On  Aug.  9, 
1921,  Mr.  Harper  stalked  another,  just  north  of  MacClenny  near  the  St.  Mary's 
River  on  an  oak  ridge.  In  both  instances  a  log  rests  over  the  entrance.  This  is 
a  favorite  cover  for  both  frogs  and  turtles.  The  turtles  seem  at  times  to  prefer 
to  dig  their  hole  under  fallen  logs  and  the  frogs  often  apparently  prefer  such 
gopher-turtle  burrows  or  burrows  with  saw  palmettos  near  by. 

July  17, 1921.  About  a  dozen  began  croaking  in  a  cypress  pond  on  Chesser fs 
Island,  Ga.,  before  darkness  came  on.  When  they  began,  one  of  the  residents 
called  it  "the  monster."  At  first  while  wading  in  the  deeper  water  of  the  pond 
in  the  early  evening,  we  would  hear  this  mysterious  note  among  the  cypress 
trees,  usually  at  their  bases  or  on  logs.  The  frog  would  espy  us  first,  and  all 
we  could  determine  was  a  big  splash,  doubtless  made  by  a  large  frog.  Finally 
we  heard  one  in  shallow  water  in  a  tussock  of  sedge  2-3  feet  across  and  i  foot 
high,  and  another  near  the  base  of  a  cypress  in  deeper  water.  After  2  hours' 
work  on  Hyla  gratiosa  we  sought  out  a  group  of  gopher  frogs  we  were  hearing 
in  the  shallower  western  portion  of  the  pond.  One  was  beside  a  cypress  tree 
in  a  depression  of  fibrous  roots.  After  we  had  focused  on  it,  it  literally  crawled 
around  to  a  different  position.  I  put  the  light  on  its  nose  and  tried  to  push  it 
back  with  my  fingers.  Then  it  leaped.  At  the  base  of  a  pine  on  a  pile  of  chips 
i%  feet  above  the  surface  was  another  croaker.  After  a  time,  it  would  "bat" 
its  eye  toward  the  bull's-eye.  We  secured  two  flashlight  photographs  of  it.  These 


FROGS:  RANIDAE  439 

frogs  were  not  hard  to  catch  when  discovered,  but  they  were  shy.  Often  when 
first  put  under  light  they  would  sink  back  in  the  fibrous  roots,  depressions, 
and  covers.  One  was  spread  out  in  the  water  among  the  spice  bushes.  The 
influence  of  near-by  croakers  seemed  to  stimulate  this  one  as  it  does  others. 
We  caught  several,  put  them  in  a  bag,  and  induced  these  to  croak.  The 
sprawled-out  specimen  responded  beautifully  with  his  croaks. 

August  17,  1922,  Trail  Ridge  near  Milliard,  Fla.  (an  oak  ridge  locality). 
After  an  excessively  rainy  period,  we  were  detouring  on  the  Dixie  Highway 
when  Mr.  Harper  heard  spadefoots  in  some  shallow  pine  barren  ponds.  Later, 
in  the  same  place,  after  5  P.M.,  I  thought  I  heard  frogs  m  the  higher  open 
places  in  the  pine  forest  near  the  spadefoot  pools.  We  suspected  they  were 
gopher  frogs  and  queried  whether  they  were  in  the  ground,  at  the  gopher- 
turtle  burrows,  or  moving  to  their  breeding  places.  We  never  found  one,  and 
one  of  the  party  thought  it  a  ventnloquial  effect  from  a  near-by  cypress  pond. 
When  darkness  came  on,  this  pond  was  suddenly  resonant  with  gopher  frog 
calls  in  chorus.  These  chose  perches  similar  to  those  described  in  1921.  One 
was  at  the  base  of  a  stump  and  above  the  water.  Another  was  in  the  notch  of 
a  floating  log.  A  third  was  at  the  base  of  a  cypress.  A  fourth  was  beside  a  stump 
and  mostly  sunk  in  the  water.  With  all  four  of  us  searching  we  saw  only  one 
croak;  the  rest  never  responded  when  we  were  near. 

Authorities'  corner: 

E.  Loennberg,  Fla.,  1895,  p.  339.  T.  Hallinan,  Fla.,  1923,  pp.  19-20. 

T.  Barbour,  Fla.,  1920,  p.  55.  A.  F.  Carr,  Jr.,  Fla.,  i94ob,  p.  64. 

Slater's  Frog,  Slater's  Spotted  Frog,  Cascade  Frog 

Rana  cascadae  Slater.  Plate  XCIII;  Map  27. 

Range  and  Habitat:  "Type — C.  P.  S.  No.  2883,  an  adult  female  from  lilysian 
Fields,  Rainier  National  Park,  Washington.  Elevation  5700  feet.  Collected  by 
the  author  June  19, 1938.  On  life  zones  Rana  cascadae  is  typically  a  Hudsonian 
frog.  I  already  have  specimens  in  my  collection  from  the  high  parts  of  the 
following  counties  of  the  State  of  Washington:  Chclan,  Clallnm,  Kittitas, 
Lewis,  Mason,  Pierce,  Skagit,  Skamania,  Snohomish  and  Yakima;  and  one 
specimen  from  Potsville,  Idaho.  It  will  likely  be  found  in  Oregon,  British 
Columbia,  and  possibly  Alaska"  (J.  R,  Slater,  Wash.,  19 ^gb,  pp.  146-149). 

In  1942  Slater  had  it  from  thirteen  instead  of  nine  (1939)  counties  m  Wash- 
ington. 

Size:  Adults,  2-3  inches.  Males  50-58  mm.  Females,  52-74  mm.  These  meas- 
urements were  taken  from  21  males  and  20  females  in  Slater's  laboratory.  The 
average  of  males  was  53.3  mm.,  the  mean  53  mm.;  the  average  of  the  females 
was  63  mm.,  the  mean  60  or  62  mm.— quite  a  difference  in  size.  The  frogs 
were  taken  April  to  October— April  10,  May  15,  30,  June  18,  July  4,  Aug.  29, 

Oct.  2, 


440  HANDBOOK  OF  FROGS  AND  TOADS 

General  appearance:  A  medium-sized  frog,  somewhat  similar  to  R.  pretiosa. 
"General  coloration  dorsal  surface  slight  greenish-brown  with  many  black 
spots  about  three  millimeters  in  diameter;  dorsolateral  folds  light  brown; 
sides  green-brown  mottled  with  black  and  fading  to  light  yellow  toward  ven- 
tral surface;  space  behind  eye  including  ear,  and  line  under  canthus  rostralis, 
dark  brown;  ridge  along  upper  jaw,  yellowish;  lips,  mottled;  throat  and 
undersurfaces  of  body  and  limbs,  very  light  yellow;  upper  surface  fore  limb, 
mottled  and  slight  bar;  hind  limb  with  broken  dark  bands"  (J.  R.  Slater, 
Wash.,  iQ3gb,  p.  146). 

Color:  Tipso  Lake,  Pierce  Co.,  Wash.,  from  J.  R.  Slater,  June  7,  1941. 
Male.  Dorsal  color  and  upper  side  of  legs  Isabella  color.  On  top  of  head  and 
between  costal  folds  and  down  to  belly  color  are  numerous  irregular  black 
or  brownish  olive  spots,  the  largest  5  mm.  across.  These  spots  are  in  five  or 
six  bars  on  rear  and  front  of  tibia,  being  broken  on  middorsal  tibial  surface. 
There  are  three  or  four  such  indistinct  bars  on  rear  of  tarsus  and  outer  toe. 
There  are  two  bars  on  antebrachium  and  a  prominent  vertical  bar  from  in- 
sertion of  brachium  halfway  to  radius  on  front  of  brachium.  The  mask  is 
brownish  olive  or  Prout's  brown,  and  from  rear  of  mouth  involves  tympanum, 
extends  forward  as  a  prominent  line  past  the  nostril  bordered  below  with  a 
prominent  cream-buff  line  from  snout  to  angle  of  mouth,  and  incurves  slightly 
along  the  infralabial  margin  of  the  rictus.  The  infralabial  margin  and  throat 
are  vermiculated  with  drab  and  ventral  color.  The  throat  is  colonial  buff,  be- 
coming deep  colonial  buff  on  belly  and  honey  yellow  on  underside  of  legs. 
The  rear  of  femur  is  Isabella  color.  Lower  part  of  eye  black  with  wash  of  drab 
and  flecks  of  chamois.  The  lower  portion  of  ins  rim  is  chamois,  but  the  upper 
part  expands  as  a  solid  spot  of  reed  yellow. 

Female.  Much  the  same  as  male  but  no  spots  or  very  few  spots  between 
dorsolateral  folds,  below  which  are  a  few  scattered  spots.  The  dorsum  is  huffy 
olive.  The  throat  is  clear  and  a  bit  yellow,  perhaps  amber  yellow.  The  legs 
are  much  the  same.  The  supralabial  fold  is  primrose  yellow  rather  sharply 
bordered  below  by  brownish  olive. 

Male.  Deer  Lake,  Sol  Due  River,  Clallam  Co.,  Wash.,  north  side  Olympic 
Mts.,  from  James  R.  Slater,  June  7,  1930.  Color  of  back  a  dark  greenish  olive 
with  several  rounded  inky  spots  on  dorsum  and  some  below  dorsolateral  folds. 
Sides  reed  yellow  to  citron  green  with  heavy  black  mottling.  Line  on  upper 
jaw  reed  yellow  to  primrose  yellow.  Under  surfaces  primrose  yellow,  with 
some  olive-ocher  on  under  surfaces  of  legs.  Legs  with  black  bars.  Mask  is 
brownish  olive.  Eye  as  in  the  female. 

Female.  Back  Saccardo's  olive,  mask  Sanford's  brown,  dorsolateral  folds 
conspicuous.  Few  to  many  black  spots  on  dorsum  and  many  along  the  sides 
on  a  pale  yellow-green  ground.  The  pale  yellow-green  conspicuous  in  axil  and 
in  groin.  Line  along  upper  jaw  is  pale  chalcedony  yellow.  Ventral  surfaces 
sulphur  yellow.  There  is  a  touch  of  salmon-orange  or  orange-buff  along 


FROGS:  RANIDAE 


441 


underside  of  tarsus.  Under  surface  of  femur  chamois  to  olive-buff.  The  iris  is 
black  with  band  of  pinard  yellow  specks  above  and  bordering  pupil  and  light 
orange-yellow  specks  below.  Around  the  outer  edge  is  a  broken  band  of  light 
lumiere  green, 

Structure:  "Form  moderately  slender;  head  broad;  external  nares  halfway 
between  tip  of  muzzle  and  orbit;  canthus  rostralis  prominent;  orbit  moderate; 
interorbital  space  about  two-thirds  length  of  orbit;  moderate  ridge  on  upper 
jaw,  beginning  anterior  to  the  orbit  and  extending  posteriorly  to  fore  limb; 
tympanum  inconspicuous,  round,  diameter  about  one-half  length  of  orbit, 
separated  from  orbit  by  about  two-thirds  of  its  diameter,  foreleg  stout;  palm 
with  three  elongated  tubercles;  sesamoid  tubercles  very  prominent;  digits 
stout,  in  order  of  decreasing  length,  3,  4,  i,  2;  dorsolateral  folds  broad  and 
prominent;  hind  limb  slim;  tibia  nearly  the  same  length  as  femur;  tarsus 
greater  than  one-third  length  of  foot;  two  metatarsal  tubercles;  toes  stout,  in 
order  of  decreasing  length,  4,  3,  5,  2,  i;  web  moderate. 

"Tongue  narrow  oval,  posterior  end  with  well  developed  lobes,  attached 
by  anterior  one-half  to  floor  of  mouth;  internal  nares  slightly  overlapped  by 
upper  jaw;  vomerine  teeth  mostly  posterior  to  line  joining  centers  of  internal 
nares,  two  areas  of  which  converge  posteriorly  and  nearly  join;  maxillary 
and  vomerine  teeth  small. 

"Surfaces  generally  smooth  with  slight  roughening  on  dorsal  surface  of 
body,  posterior  femur,  and  dorsal  tibia"  (J.  R.  Slater,  Wash.,  i9J9b,  p.  145). 

Voice:  No  record.  Of  the  frogs  from  Tipso  Lake  (Pierce  Co.,  Wash.),  June 
7,  1941,  sent  me  by  J.  R.  Slater  this  note  was  made:  There  are  no  external 
vocal  sacs.  While  in  captivity,  they  have  croaked  several  times,  especially  when 
seized  ahead  of  the  hind  legs.  It  is  a  low  croak. 

Breeding:  Females  shipped  to  us  June  7,  1930,  by  J.  R.  Slater  and  collected 
at  Deer  Lake,  Sol  Due  River,  Clallam  Co.,  Wash.,  on  north  side  of  Olympic 
Mts.  Enroute  they  laid  eggs  in  the  moss  of  the  container.  These  may  not  have 
expanded  normally  but  they  appear  to  be  far  apart  like  Rana  sylvatica  eggs. 
They  seem  to  have  an  inner  envelope  with  the  vitellus  from  2.2-2.8  mm.  or 
more  in  diameter. 

"Eggs. — The  different  jelly  envelopes  average  as  follows:  outer  11.3,  middle, 
5.8,  inner  4.9,  while  the  egg  proper  is  2.25.  Color  of  egg  is  very  dark  on  upper 
three-fourths  and  light  cream  below. 

"Tadpoles. — The  teeth  are  in  two  upper  rows  with  a  small  gap  in  center 
of  second  row  and  four  lower,  three  of  which  are  nearly  full  length  while  the 
fourth  is  about  one-fourth  the  length  of  the  others.  Tadpoles  at  hindlimb 
bud  stage  are  34  millimeters  and  when  the  toe  buds  show  on  hind  limb  they 
are  about  40  millimeters. 

"Life-history. — Spawning  period  is  from  May  20  to  July  10  depending  on 
how  much  snow  there  is  to  melt  before  the  ponds  appear;  eggs  deposited  in 
small  ponds  in  numbers  up  to  425  by  single  female,  hatch  in  8  to  20  days; 


442  HANDBOOK  OF  FROGS  AND  TOADS 

larval  period  80  to  95  days,  with  very  few  tadpoles  living  over  to  following 
summer;  transformation  size  for  most  of  them  14  mm.  or  less;  size  at  one 
year  after  transforming  about  30  mm.;  sexual  maturity  reached  at  end  of  three 
full  years  or  possibly  four"  (J.  R.  Slater,  Wash.,  i939b,  p.  149). 

Journal  notes:  The  last  of  March,  1942,  we  sought  to  see  this  frog,  but  Pro- 
fessor Slater  told  us  that  we  were  too  early  for  this  form  of  the  higher  eleva- 
tions. 

March  3i-April  2.  Worked  over  Slater's  R.  cascadae;  measured  for  range 
of  adults  male  and  female  and  transformation.  He  says  R.  cascadae  has  no  red 
underneath  but  is  cream  or  straw  color.  Most  of  his  records  are  above  4000  ft. 
He  contrasts  it  with  R.  aurora  aurora.  He  showed  us  two  skins  of  R.  a.  aurora 
and  R.  pretwsa  to  show  differences  in  coloration  and  color.  Aurora  is  mottled 
black  and  yellow  with  some  green  in  groin;  pretiosa  is  red.  Does  R.  cascadae 
run  into  R.  b.  boyln  in  Cascades?  He  has  a  number  from  Breitenbush  Lake, 
Marion  Co.,  Ore.  Slater  conceives  of  R.  cascadae  as  closest  to  R.  a.  aurora.  It  is 
different  from  R.  b.  boyln.  It  is  also  different  from  R.  p.  pretiosa.  What  is  its 
relation  to  jR.  p.  lutewentm?  (See  R.  a.  aurora).  Cursory  notes  on  Slater's 
material  follow: 

1.  He  has  many  interesting  young  specimens.  One  lot  just  transformed 
measures  13,  13,  14,  15,  16,  17  mm.  The  tadpoles  belong  in  wood  frog  group, 
a  R.  temporana  type.  Longest  tadpole  is  70  mm.,  of  which  body  is  22  mm., 
depth  of  body  14%  mm.,  and  depth  of  tail  crest  15  mm.  The  tail  crest  comes 
forward  onto  body  almost  to  the  vertical  of  spiracle;  spiracle  is  nearer  vent 
than  tip  of  snout;  eye  is  entirely  dorsal.  Another  tadpole  is  70  mm. 

2.  June  1 8,  1938.  Two  23  mm.  specimens  just  transformed;  one  young 
28  mm. 

3.  June  18,  1938.  Among  the  adults  are  three  young,  one  24  mm.  and  two 
36  and  38  mm. 

4.  July   n,  1931.  One  transformed  at  20  mm.;   intermediate  specimen 
43  mm. 

5.  Aug.  5,  1937.  Several  transformed  23.5,  23.5  mm.;  past  transformation 
25, 34  mm. 

6.  Aug.  16,  1930.  One  21  mm.  just  transformed;  one  32  mm. 

7.  Oct.  10, 1939.  One  just  transformed  20.6  mm.,  another  22.6  mm.,  and  one 
36  mm. 

8.  We  noted  one  specimen  marked  "male  not  strong"  at  38  mm.;  and  one 
"female  not  mature"  at  44  mm. 

9.  Random  notes  on  specimens  measured  follow: 

a.  Young,  28  mm.,  has  2  prominent  metatarsal  tubercles. 

b.  Young,  36  mm.,  has  2  prominent  metatarsal  tubercles. 

c.  Young,  36  mm.,  heel  goes  to  snout's  tip. 

d.  Female,  44  mm.,  not  mature.  Hardly  spotted  at  all;  quite  a  few  tubercles 


FROGS:  RANIDAE 


443 


Plate   XCllI.    Rana   cascadae    (X%)- 
1,2,3,5.  Males.  4,6.  Females. 


Plate  XCIV.  Rana  catesbeiana,  1,4,5. 
Males  (X%).  2-  Female  (X%).  3-  Tad- 
pole  (X%).  6.  Eggs  (X%).  - 


444  HANDBOOK  OF  FROGS  AND  TOADS 

on  back  and  side.  Heel  goes  to  nostril;  there  is  a  dark  spot  wherever  there  is 
a  tubercle  which  serves  as  a  light  center. 

e.  Males,  56  mm.,  56  mm.;  female,  56  mm.  One  male  very  spotted;  one 
male  with  heel  almost  to  tip  of  snout.  Male  has  web  from  tip  of  fourth  toe 
convex  and  folded  in  one  place,  the  web  from  fourth  to  third  somewhat  con- 
vex, from  third  to  second  about  straight,  but  hind  foot  webbing  is  much  more 
pronounced  than  in  the  female.  In  the  female  the  web  is  concave  between 
all  toes.  Tympanum  same  size  but  more  oblique  in  male  than  m  female. 
Thumb  of  male  not  as  much  bilobed  as  in  some  western  frogs.  All  fingers  of 
male  are  much  thicker,  also  brachium  and  antebrachium. 

f.  Female,  68  mm.  Internasal  space  broad,  flat;  interorbital  space  rather 
narrow  and  less  than  internasal;  uppei  eyelid  rather  broader  than  interorbital 
space;  dorsolateral  folds  rather  flat  but  prominent,  broader  in  the  forward  half. 
Skin  between  dorsolateral  folds  smooth,  below  folds  quite  a  few  tubercles; 
fore  and  hind  limbs  almost  smooth  above;  dorsolateral  fold  starts  below  eye 
and  then  just  below  the  tympanum  doubles  in  width.  It  is  light  in  color  and 
this  makes  a  prominent  light  wart  back  of  corner  of  mouth.  Rear  of  femur 
uniform.  Two  metatarsal  tubercles,  inner  (opposite  toe  5)   is  very  slight. 
Prominent  palmar  tubercle  at  base  of  third  and  fourth  digits.  Tongue  promi- 
nently or  deeply  notched.  Vomerine  patches  between  choanae  almost  meet 
on  the  meson.  Articular  tubercles  on  foot  not  very  prominent.  Specimen  not 
much  spotted. 

Authorities9  corner:  There  is  as  yet  only  one  authority  for  this  species,  Prof. 
J.  R.  Slater,  who  kindly  showed  me  all  his  material. 

"Some  previous  collectors  have  placed  specimens  of  this  species  with  Rana 
aurora  aurora  while  others  have  called  them  Rana  pretiosa  but  it  appears  to 
me  that  Rana  cascadae  may  be  more  primitive  than  these  other  two  species. 
This  species  is  named  from  the  region  in  which  I  first  found  it  which  is  the 
Cascade  Range"  (J.  R.  Slater,  Wash.,  1939^  p.  149). 

Bullfrog,  Bloody  Nouns,  Bully,  Jug-o'-Rum,  North  American  Bullfrog, 

American  Bullfrog 

Rana  catesbeiana  Shaw.  Plate  XCIV;  Map  29. 

Range:  From  Laredo  or  the  mouth  of  the  Pecos  River  through  the  Pan- 
handles of  Texas  and  Oklahoma,  extreme  western  Kansas,  Nebraska  to 
Minnesota,  Lake  Huron  to  Maine.  Absent  from  southern  half  of  Florida. 
Introduced  in  New  Mexico,  Colorado,  Wyoming,  Idaho,  Arizona,  Nevada, 
California,  Oregon,  Washington,  and  British  Columbia. 

Habitat:  Strictly  aquatic,  these  frogs  seem  to  prefer  millponds,  hydraulic 
lakes,  reservoirs,  and  kindred  bodies  of  water. 

The  authors'  best  collecting  grounds  have  been  a  clear  glacial  lake  in  a  New 


FROGS:  RANIDAE  445 

England  kettle  hole,  with  a  slight  suggestion  of  the  sphagnous  flora  about 
it;  a  pond  in  a  clear  trout  brook;  a  large  reservoir  for  a  hydraulic  laboratory; 
a  disused  millpond;  and  a  wooded  lake  whose  shifting  water  level  had  made 
a  fringe  of  overhanging  dead  trees,  floating  logs,  and  submerged  roots  and 
limbs.  In  every  case  the  shores  were  more  or  less  wooded,  but  more  important 
are  two  factors:  (i)  shallows  where  the  species  can  transform,  and  (2)  brush, 
stumps  whose  roots  are  at  the  edge  of  the  pond,  driftwood  along  the  bank, 
or  matted  roots  of  a  fringe  of  willow  trees. 

When  croaking  begins,  the  males  often  take  certain  perches  in  which  they 
keep  a  proprietary  interest.  About  one  pond  the  authors  once  located  seven 
such  places,  each  with  its  possessor,  only  once  finding  two  in  one  place.  The 
first  frog  was  on  a  board  in  water  filled  with  brush;  another  was  perched 
on  a  log  among  brush  beneath  the  float  of  a  boathouse;  the  third  was  on  the 
bank  among  some  limbs  extending  into  the  water;  the  fourth  was  by  an 
overturned  stump  whose  roots  were  partly  submerged;  the  fifth  was  among 
driftwood  along  the  shore;  the  sixth  was  on  a  stationary  float;  and  the  last 
was  at  the  base  of  a  tree  fallen  into  the  pond.  When,  however,  the  males 
about  a  lake  are  numerous  enough  to  make  their  night  croaking  a  real  chorus, 
it  is  not  likely  that  any  one  individual  holds  a  favorite  site  to  the  exclusion 
of  the  others.  The  bullfrog  is  a  solitary  form. 

For  migration  and  growth  sec  E.  C.  Raney,  N.Y.,  1940,  pp.  733-745. 

"Moderately  common;  frequents  margins  of  ponds,  ditches  and  swamps" 
(O.  C.  Van  Hyning,  Fla.,  1933,  p.  4). 

Size:  Adults,  5%-8  inches.  Males,  85-180  mm.  Females,  89-184,  possibly 
200  mm.  The  species  seldom  breeds  before  it  reaches  4  inches  (100  mm.). 

General  appearance:  The  bullfrog  is  our  largest  frog.  It  is  rather  broad  in 
body.  In  the  North  it  is  usually  a  greenish  drab  on  the  back  and  yellowish 
white  underneath.  The  skin  may  be  roughened  with  fine  tubercles.  There  is 
no  lateral  fold,  except  a  short  fold  over  and  behind  the  tympanum.  There 
are  a  few  dusky  spots  on  the  legs.  This  frog  in  the  Gulf  region  may  be  so 
dark  as  to  be  almost  black  on  the  back  and  heavily  mottled  beneath,  and  even 
the  webs  of  the  feet  may  be  almost  black. 

Color:  Female.  Chopowampsie  Swamp,  Va.,  April  22,  1929.  Head,  before, 
below,  and  back  of  eyes,  upper  eyelids,  and  area  from  angle  of  mouth  to 
tympanum  are  cosse  green,  callistc  green,  or  courge  green.  Rest  of  upper  parts 
same  color  or  light  yellowish  olive,  yellowish  citrine,  or  oil  yellow  incon- 
spicuously spotted  with  Saccardo's  olive  or  light  brownish  olive.  Rim  on  outer 
part  of  tympanum  same  color.  Hind  and  tore  limbs  same  color  as  back,  or 
ecru-olive  with  yellowish  olive  or  dark  greenish  olive  crossbars  on  femur  and 
tibia  and  same  color  in  spots  on  fore  limbs.  Front  of  femur  reticulated  with 
grayish  olive,  also  groin  and  undersides  of  hind  legs  partially  reticulated  or 
spotted  with  light  grayish  olive  with  white  interspaces.  Under  parts  white  or 


446  HANDBOOK  OF  FROGS  AND  TOADS 

pale  olive-buff.  Ins  black  spotted  with  rufous,  apricot  orange,  or  orange 
chrome.  Pupil  rim  pale  lemon  yellow. 

Structure:  No  ridges  down  either  side  of  back;  web  of  fourth  toe  extends 
to  its  tip;  male  with  internal  vocal  sacs,  when  sacs  inflated  a  flattened  pouch 
beneath  chin;  male  with  tympana  larger  than  eye;  head  narrow. 

Voice:  The  sonorous  bass  notes  have  received  countless  characterizations, 
among  which  are  the  familiar  jug-o'-rtim,  more  rum,  blood  'n'  ouns,  br~wiim, 
be  drowned,  \nee  deep,  and  bottle-o'-rum.  These  notes  have  wonderful  carry- 
ing power  and  are  commonly  heard  in  the  evenings  of  early  summer. 

Breeding:  In  the  North  bullfrogs  breed  the  last  of  June  or  in  July  when  the 
air  temperature  is  about  80°  and  the  water  has  warmed  up  to  70°.  In  the 
South  they  breed  much  earlier.  We  found  an  egg  mass  in  San  Antonio,  Tex., 
Feb.  12, 1925.  The  egg  mass  is  a  large  surface  film  2  by  %,  2  by  2,  or  2  by  2-J4 
feet;  the  eggs,  black  and  white,  are  /{>o-Vi;>  inch  (1.2-1.7  mm.)  in  diameter. 
There  is  no  inner  envelope  and  the  outer  merges  into  the  jelly  film.  The  tad- 
pole is  large,  4-6%  inches  (100-165  mm.),  olive  in  color  with  fine  speckings 
of  black,  and  with  the  tooth  ridges  %,  rarely  %.  In  the  North  it  spends  two 
winters  as  a  tadpole,  and  transforms  from  July  i  to  Aug.  15,  or  in  the  South, 
from  last  of  May  onward,  at  1^/4-2%  inches  (31-59  mm.). 

See  L.  R.  Aronson,  N.Y.,  1943,  pp.  4-6. 

Journal  notes:  McLean,  N.Y.  On  June  16,  191?,  when  we  discovered  some 
seven  or  eight  egg  complements,  considerably  more  light  was  thrown  on  the 
egg  laying.  More  than  ever  arc  we  of  the  opinion  that  in  the  expulsion  of  the 
eggs  the  two  forms,  green  frog  and  bullfrog,  agree.  In  several  of  the  comple- 
ments the  egg  masses  were  in  brush  near  the  bank's  edge,  and  all  of  these 
either  were  of  the  film  type  or  had  originally  been  so.  In  one  case  the  eggs 
had  the  spoiled  appearance  which  green  frogs'  eggs  sometimes  have  when 
they  have  been  stranded  high  and  dry.  The  fresh  films,  1%  feet  or  more  in 
diameter,  we  could  easily  detect  because  of  the  conspicuous  air  bubbles  in  the 
mass. 

Along  the  east  edge  of  this  particular  lake  occurs  the  deepest  water,  and 
here  for  many  years  the  tadpoles  have  often  transformed.  We  could  not  under- 
stand this  deep-water  transformation  site.  Frequently  the  bullfrogs  laid  their 
films  in  midpond  around  stumps,  or  perchance  attached  the  surface  egg  film 
to  the  tips  of  some  dead  elderberry  stems  reaching  8  feet  out  into  the.  water. 
The  strong  western  winds  create  waves  on  the  water's  surface,  breaking  up 
the  disks  into  small  packets.  These  drifted  to  the  east  side  and  we  often  de- 
tected them  there  among  the  algae,  their  position  revealed  by  the  foamy  air 
bubbles.  At  this  east  edge  they  hatch,  remain  in  the  deep  water,  and  hence 
transform  here  instead  of  in  the  usual  transforming  site,  the  shallows. 

June  30, 1913,  Otter  Lake,  Dorset,  Ont.  Evening.  Bullfrogs  in  chorus.  They 
are  common  here.  In  the  grassy  run  west  of  Brennan's  house,  they  are  common 
and  lie  on  the  surface  of  the  water.  If  frightened  they  dive  and  soon  come 


FROGS:  RANIDAE  447 

up  again  a  short  distance  away.  The  other  day  the  boys  caught  29  with  a  red 
flannel  hook.  The  following  evening  with  electric  flashlight  we  caught  15.  The 
next  day,  June  29,  Charles  and  I  in  the  daytime  went  up  to  the  N.  W.  swampy 
corner  of  the  lake,  where  untold  logs  and  stumps  are,  and  captured  20  more. 
When  I  cleaned  them,  none  of  the  females  had  laid.  They  were  still  with  eggs. 
Furthermore,  we  found  no  eggs  in  the  field.  Surely  this  is  the  wrong  season 
to  catch  them.  Their  capture  should  not  be  allowed  until  the  middle  of  July 
or  beginning  of  August. 

June  4, 1917.  Camped  at  Broad  River,  S.C.  In  long  shallow  pools  2-3  ft.  deep, 
where  plenty  of  yellow  water  lilies  grow,  found  three  masses  of  bullfrog  eggs. 
One  fresh  mass  was  21/£  by  i1/-*  ft.  in  diameter  and  was  at  the  surface  around 
the  stems  of  a  yellow  water  lily.  Another  mass  looked  like  cooked  rhubarb 
and  was  more  or  less  dirty.  In  another  case,  the  egg  mass  was  on  Cham  which 
surrounded  the  stems  of  the  water  lilies.  Here  there  were  plenty  of  adult  bull- 
frogs. 

Feb.  12,  1925,  Brackenridge  Park,  San  Antonio,  Tex.  Near  a  ford  below  the 
new  swimming  pool  found  two  large  films  of  eggs.  Instinctively  I  called  them 
R.  catesbeiana  but  why  February?  The  water  was  at  least  55°  to  60°  at 
3:30  P.M.  The  air  according  to  the  San  Antonio  weather  bureau  was  65°  to  64°. 
One  egg  mass  was  2M*~3  ft.  across.  Film  among  algae  and  MyriophyUum-lilic 
plant.  The  inner  edge  of  mass  about  one  foot  from  bank  in  water  4-6  in.  deep. 
A  little  below  a  camper  said  he  had  heard  the  bullfrogs  croak,  and  in  this 
locality  were  the  large  tadpoles  of  R.  catesbeiana.  Then  at  the  place  where  the 
mass  was,  there  were  two  bullfrogs  with  heads  up,  one  a  half-grown  one  and 
one  a  very  large  one. 

Introduction  west  of  Rocf{y  Mountains.  Several  young  naturalists  have  sent 
us  strange  frogs  from  the  West.  "Are  they  new?"  "They  do  not  agree  with 
the  descriptions  of  any  frog  of  the  Plateau  or  Pacific  States."  "What  arc  they?" 
Usually  the  perplexing  form  is  a  young  or  half-grown  introduced  bullfrog. 

To  us  as  naturalists  the  presence  in  the  extreme  West  of  the  bullfrog  is  un- 
desirable. According  to  the  following  authorities  this  species  has  been  intro- 
duced into  Colorado  (Ellis  and  Henderson),  New  Mexico  and  Arizona 
(Bogert),  Idaho  (Slevin),  Washington  (Slater),  British  Columbia  (Carl), 
Oregon  (Gordon),  Nevada  (Linsdale),  and  California  (Slevin).  Many  who 
think  of  immediate  results  might  proclaim  it  a  blessing.  It  is  a  voracious, 
omnivorous,  carnivorous  species.  We  would  hate  to  see  it  introduced  in  Myer's 
Bufo  exsul  colony.  It  is  already  in  the  restricted  R.  fisheri  habitat.  The  follow- 
ing cursory  notes  of  the  summer  of  1942  show  how  wideprcad  it  is: 

March  8,  1942,  California.  Went  to  Searsbury  Reservoir.  Heard  a  bullfrog 
there.  Saw  no  frogs. 

March  30,  Washington.  Went  on  short  trip  .  .  .  with  Mr.  James  Slater  to 
Sparaway  Lake.  There  are  no  end  of  R.  catesbeiana  in  the  lake.  In  a  man's  pool 
near  the  pond  saw  several  R.  catesbeiana. 


448  HANDBOOK  OF  FROGS  AND  TOADS 

April  8,  above  Medford,  Ore.  Stopped  on  Rogue  River  east  of  Beagle.  Here 
in  a  dug  pond  there  were  numerous  bullfrogs  leaping  into  the  water.  A  short 
distance  above  Trail  saw  a  series  of  brushy  pools  with  immense  bullfrogs. 

April  n,  Chico  to  Davis,  Calif.  Two  miles  north  of  Richvale,  in  bordering 
ditches  filled  with  cattails,  were  bullfrogs. 

May  15,  Las  Vegas,  Nev.  Went  out  toward  Tonopah,  25  miles  and  returned 
after  dark.  Near  a  trailer  camp  bullfrogs  calling  in  a  large  pond. 

May  16,  Las  Vegas.  At  the  U.S.  Fish  Hatchery  springs  found  bullfrogs  but 
no  R.  fisheri. 

May  20,  Moapa  Indian  Reservation  (near  Glendale,  Ncv.).  Bullfrogs  in 
their  drainage  ditches. 

June  15,  Arizona.  Went  out  last  night  toward  Florence  Jet.  Heard  bullfrogs 
and  Hammond's  spadefoot  in  pool  at  Desert  Wells. 

July  21,  LaPosta  Lodge,  El  Paso,  Tex.  On  road  toward  Smelter  town  took 
one  Rana  catesbeiana. 

m 

Authorities9  corner:  "It  is  the  General  Belief  of  the  People  in  Virginia  that 
they  keep  the  springs  clean,  and  purify  the  Water,  wherefore  they  never  kill 
or  molest  them,  but  supcrstitiously  believe  it  bodes  them  ill  so  to  do. 

"The  Noise  they  make  has  caused  their  Name;  for  at  a  few  Yards  Distance 
their  Bellowing  sounds  very  much  like  that  of  a  Bull  a  quarter  of  a  Mile  off, 
and  what  .adds  to  the  Force  of  the  Sound,  is  their  sitting  within  the  hollow 
Mouth  of  the  Spring"  (M.  Catesby,  1731-1743,  p.  72). 

"At  Baltimore  I  first  saw  the  fire-fly.  .  .  .  Mischievous  boys  will  sometimes 
catch  a  bull-frog,  and  fasten  them  all  over  him.  They  show  to  great  advantage, 
while  the  poor  frog,  who  cannot  understand  the  'new  lights'  that  are  breaking 
upon  him  affords  amusement  to  his  tormentors  by  hopping  about  m  a  state 
of  desperation1'  (G.  T.  Vigne,  Gen.,  1832,  p.  140). 

"The  frogs  were  as  great  a  source  of  amusement  to  us  as  the  pigeons  were 
of  excitement.  Wherever  there  was  a  spot  of  water,  the  double  bass  of  the 
bullfrogs  striking  in  every  now  and  then  amidst  indescribable  piping  of  the 
multitudes  of  their  smaller  brethren.  Il  is  very  difficult  to  catch  a  sight  of  these 
bassoon  performers,  as  they  spring  into  the  water  at  the  slightest  approach 
of  danger;  yet  you  may  now  and  then  come  on  them  basking  at  the  side  of 
a  pond  or  streamlet,  their  great  goggle  eyes  and  black  skin  making  them 
look  very  grotesque.  They  are  great  thieves  m  their  own  element,  many  a 
duckling  vanishing  from  its  mother's  side  by  a  sudden  snap  of  some  one  of 
these  solemn  gentlemen  below.  They  are  a  hungry  race,  always  ready  appar- 
ently for  what  they  can  get,  and  making  short  work  with  small  fishes,  all 
kinds  of  small  reptiles,  and  even,  I  believe,  the  lesser  kinds  of  snakes,  when 
they  can  get  them.  These  fellows  are  the  giants  of  the  frog  tribes,  and  portly 
gentlemen  withal,  some  of  them  weighing  very  nearly  a  pound"  (J.  C.  Geikie, 
Gen.,  i882(?),  pp.  216-217). 


450  HANDBOOK  OF  FROGS  AND  TOADS 

Green  Frog,  Pond  Frog,  Spring  Frog,  Common  Spring  Frog,  Bullfrog, 
Bawling  Frog,  Yellow-throated  Green  Frog,  Belly  Bumper,  Bully 

Rana  damitans  Latreille.  Plate  XCV;  Map  30. 

Range:  Southeastern  Texas  to  Louisiana,  Arkansas,  southeastern  Okla- 
homa, eastern  Kansas  through  Iowa,  western  Minnesota  to  Canada  east  to 
Gaspe  Peninsula.  Not  in  southern  half  of  Florida. 

Habitat:  A  solitary  species.  Like  the  bullfrog,  it  lives  in  swamps  and  in  our 
deeper,  larger  ponds  and  reservoirs,  but  the  green  frog  also  lives  in  smaller 
ponds  and  pools.  In  fact,  along  watercourses  there  is  hardly  a  small  pond 
which  cannot  claim  a  green  frog.  Hibernates  in  the  water. 

"Bayhcads  and  titi  swamps;  wet  low  hammocks,  streams  in  mesophytic 
hammock;  cypress  swamps.  .  .  .  Not  common  .  .  ."  (A.  F.  Carr,  Jr.,  Fla., 


Size:  Adults,  2-4  inches.  Smaller  in  the  South.  Males,  52-72  mm.  in  the 
South,  and  reaching  95  mm.  in  the  North.  Females,  58-75  mm.  in  the  South, 
reaching  100  mm.  in  the  North. 

General  appearance:  This  is  a  yellow-throated  bullfrog.  In  the  North  the 
green  frog  is  one  of  our  largest  frogs.  It  has  a  prominent  dorsolateral  fold 
with  a  branch  extending  almost  at  right  angles  downward  toward  the  tympa- 
num, which  is  large  and  conspicuous.  The  general  color  is  a  greenish  brown 
with  a  bright  green  mask  from  the  tympanum  forward  along  the  jaw.  The 
skin  may  be  slightly  roughened  with  small  tubercles.  The  legs  have  dusky 
bars.  An  occasional  green  frog  has  a  few  scattered  black  spots  on  the  back. 
They  are  white  beneath,  the  male  having  a  yellow  throat. 

Color:  Male.  West  wood,  N.J.,  from  A.  M.  and  C.  C.  Van  Deman,  April 
30,  1928.  Upper  parts  forward  ramctte  green  or  parrot  green.  Below  eye  and 
forward,  callistc  green.  Rear  upper  paits  olive-citrine.  Upper  parts  of  fore  and 
hind  legs  citrine  or  buffy-citrine.  Tympanum  Saccardo's  olive,  with  center 
yellowish  citrine.  Barred  with  black  on  hind  legs  and  feet.  Side  of  body  has  sea- 
foam  yellow  interspaces  between  citrine-drab  areas.  Throat  barium  yellow. 
Rest  of  under  parts  white,  groin  with  cream  color  cast.  Under  parts  of  hind 
legs  dark  vinaceous-brown.  Front  of  hind  legs  with  a  line  of  Hay's  brown. 
Line  under  tympanum  neva  green,  becoming  vinchnc  yellow  over  shoulder. 
Line  on  upper  jaw  and  lower  jaw  medal  bronze.  Front  part  of  lateral  fold 
blackish,  a  blackish  bar  back  of  tympanum,  a  raw  umber  line  on  front  of 
foreleg.  Ins  black,  pupil  rim  greenish  yellow.  Iris  above  and  below  pupil 
with  much  xanthme  orange. 

Female.  Forward  upper  parts  parrot  green,  or  cedar  green  ahead  of  eyes; 
spinach  green  below  eye  and  under  tympanum.  Rear  parts  Saccardo's  olive, 
also  upper  part  of  fore  and  hind  legs.  Tympanum  Saccardo's  olive,  center 
mummy  brown.  Top  of  hind  legs  with  black  spots  and  transverse  bars.  Rear 
of  femur  uniform  Prout's  brown  or  raw  umber.  Sides  of  body  pale  cendre 


FROGS:  RANIDAE 


451 


green  and  black.  Under  parts  white,  heavily  spotted  with  black  except  white 
throat.  Lower  jaw  edge  dotted  white  and  olive.  Iris  same  as  in  male. 

Structure:  A  large  frog;  skin  smooth;  broad  short  fold  of  skin  on  which  is 
a  yellowish  white  stripe  extending  from  angle  of  mouth  over  arm  insertion; 
dorsolateral  fold  /{>-%  distance  to  hind  limb;  fourth  toe  free  of  webbing  on 
last  two  phalanges;  male  with  enlarged  thumb  and  tympanum;  male  with 
ring  of  yellow  in  tympanum;  male  with  throat  a  bright  yellow;  the  swollen 


R.  p.  prefiosa 


Map  jo 

throat  of  a  croaking  male  is  a  flattened  pouch,  not  a  ball-like  sac;  the  vocal 
sacs  are  internal. 

Voice:  The  call  is  low-pitched,  explosive.  The  note  resembles  the  sound 
made  by  plucking  the  string  of  a  bass  viol  or  the  twang  of  a  rubber  band 
slightly  stretched  over  an  open  box.  The  male  rests  on  a  mat  of  vegetation  or 
among  grass  or  aquatic  plants,  or  freely  sprawls  out  on  the  water.  Usually 
the  hind  quarters  are  slightly  submerged. 

Breeding:  In  the  North  they  breed  from  the  end  of  May  to  mid-August, 
but  in  the  South  they  are  late  breeders.  The  black  and  white  eggs  are  in  a 
surface  film  usually  less  than  a  foot  square  and  number  iooo~4<x>o.  The  egg 
is  Vi«J  inch  (1.5  mm.),  the  inner  envelope  elliptic,  the  outer  merging  into 
the  jelly  film.  The  olive  green  tadpole  is  large,  2%*  inches  (64  mm.),  its  tail 
elongate,  green  mottled  with  brown,  and  with  tip  acute.  The  tooth  ridges 


452  HANDBOOK  OF  FROGS  AND  TOADS 

are  %,  rarely  %•  It  spends  one  winter  as  a  tadpole  and  transforms  from 
April  to  September  at  %-!%  inches  (23-38  mm.).  Many  transform  at  a 
smaller  size  in  the  South.  We  wonder  whether  the  foregoing  egg  and  tad- 
pole measurements  obtain  with  the  smaller,  browner,  southern  specimens  of 
this  species.  Do  they  always  winter  over  as  tadpoles  in  the  South?  Does  this 
long-extended  summer  breeder  of  the  North  have  a  long  season  of  ovula- 
tion  in  the  South? 

Journal  notes:  June  21,  1907,  Ithaca,  N.Y.  At  the  surface  in  the  middle  of 
the  northeast  slaughter  house  pond  we  found  at  12:30  P.M.  eggs  laid  by  six 
pairs.  In  one  case  the  complement  was  distributed  in  four  groups  about  a 
foot  apart.  In  other  cases  they  were  in  one  mass.  They  were  readily  distin- 
guishable because  of  an  excessive  amount  of  air  bubbles  among  them. 

June  1 6,  1910,  Ithaca,  N.Y.,  5:10  P.M.,  air  82°.  Heard  Rana  clamttans.  In 
west  stretch  found  one  solid  mat  of  waterweed.  On  top  of  this  at  surface  a 
film  9  inches  in  diameter  apparently  fresh.  Water  right  at  spot  82°.  Little 
farther  on  two  other  films,  irregular  crescents,  one  on  open  water.  In  shallow 
water,  two  bunches  2  days  old,  one  other  bunch  a  composite  of  two,  with 
diameter  15  by  10  inches.  Another  bunch  in  deep  water,  fresh  but  already 
distributed  in  little  packets.  Along  the  west  side  five  fresh  films,  one  14  by 
9  inches,  but  looks  like  two.  Another  bunch  about  hatched.  Another  on  Chara 
at  the  surfaces  as  usual,  a  striking  foamy  mass. 

Hellgate  Ponds,  Adirondacks,  Onekio,  N.Y.,  June  13,  1923.  Pratt's  Manual 
uses  the  character  of  spots  or  no  spots  on  the  back  to  separate  the  mink  frog 
from  the  green  frog.  Imagine  our  surprise  in  finding  practically  all  the  green 
frogs  of  these  parts  distinctly  spotted!  Green  frogs  may  have  a  few  scattered 
dark  spots,  but  not  with  the  spotting  all  over  the  upper  parts  like  these  individ- 
ual specimens.  Some  of  these  much-spotted  frogs  are  of  the  general  color  of 
a  green  frog,  some  are  very  hard  to  distinguish  from  mink  frogs,  and  one  is 
a  blue  frog  which  we  jokingly  called  our  blue  "green  frog."  This  one  was 
deep  glaucous  gray,  deep  bluish  glaucous,  or  greenish  glaucous-blue  with 
small  black  spots  all  over  upper  parts. 

June  16, 1924,  Ithaca,  N.Y.  About  9  A.M.  (sun  shining  brightly)  along  north 
side  of  my  pond,  a  pair  leaped  in.  In  grassy  corner  saw  spread  out  in  water 
with  head  under  water,  an  apparently  exhausted  or  dead  frog.  Started  to  pick 
it  up  and  another  beneath  it  swam  away.  The  upper  one  was  a  male.JSuspect 
the  lower  one  was  a  female.  At  9:50  came  to  same  corner  and  found  a  fresh 
mass  which  must  have  been  laid  soon  after  9.  Right  where  the  supposed  dead 
frog  was,  found  another  fresh  mass  just  laid.  Vegetative  pole  not  turned  down 
yet.  Here  I  found  a  large  male  with  a  small  male  embracing  it  in  the  axils. 
The  second  male  in  turn  was  embraced  by  another  male,  making  three  males. 
Then  the  head  of  the  first  male  was  held  by  a  fourth  male  whose  nose  touched 
the  second  male.  The  fourth  male  was  embraced  by  a  fifth  one.  The  impulse 
must  have  been  very  great  at  the  laying  of  the  one  female  a  while  before.  This 


FROGS:  RANIDAE 


453 


XCF.  Rafla  damitans.  i.  Male 
(X%).  2.  Female  (XH).  3-  Hind  foot 
(X%).  4-  Eggs  (X%).  5-  Female  (X%)- 


Ptof  XCF/.  Ram  fishen.  i.  Male  (X%)- 
2,3,4.  Females  (X%)- 


454  HANDBOOK  OF  FROGS  AND  TOADS 

is  not  exactly  sex  distinction.  About  10:15  I  returned  to  the  spot.  The  first 
and  second  males  were  the  same  only  turned  around  and  the  fourth  male 
was  still  hanging  on  to  the  head.  One  of  the  other  males,  doubtless  third  or 
fifth,  was  near  by,  but  not  in  embrace.  The  temperature  of  the  water  where 
the  eggs  are  is  70° F. 
Authorities'  corner: 
G.  H.  Loskiel,  (Jen.,  1794,  p.  89. 
A.  M.  Banta,  Ind.,  1907,  p.  22. 
J.  A.  Weber,  N.Y.,  1928,  pp.  109-110. 

Nevada  Frog 
Rana  fishen  Stejneger.  Plate  XCVI;  Map  34. 

Range:  Las  Vegas,  Nev. 

Habitat:  Spring  basins  or  trickling  streams  in  springy  fields.  Isolated  in 
such  spots  because  of  surrounding  desert. 

"Inhabits  the  springs  and  shoit  streams  in  the  near  vicinity  of  Las  Vegas, 
close  to  2000  feet  elevation.  These  frogs  arc  most  active  in  spring  and  early 
summer;  dates  of  capture  range  from  early  in  March  to  August  13"  (J.  M. 
Lmsdale,  Nev.,  1940,  p.  210). 

August  20,  1925,  Las  Vegas,  Nev.  Went  out  the  Tonopah  road  to  Tule 
Springs,  a  distance  of  about  16  miles.  The  springs  are  large  and  fine,  but 
yielded  us  only  water  for  our  radiator.  Then  we  turned  back  and  about  3  miles 
from  Las  Vegas  passed  an  artesian  well  pouring  out  a  big  stream  of  water, 
and  then  came  to  a  low  rather  swampy  area  with  quite  a  stream  rushing  on 
one  side  of  it.  No  trogs  seen  here.  We  drove  a  short  distance  to  a  place  where 
on  both  sides  of  the  road  small  springs  come  out  of  the  ground  in  basins  per- 
haps 3-4  feet  across,  and  run  as  trickling  streams  down  into  the  field.  These 
had  been  artificially  ditched  in  places.  The  streams  are  flowing  but  covered 
with  an  alga.  The  region  is  grassv  with  a  stiff  grass.  The  immediate  edges 
are  sedgy.  I  first  saw  a  frog  jump  into  a  tiny  stream  on  the  west  side,  just 
showing  through  a  break  in  the  algae.  We  caught  him.  He  is  the  game  we 
came  here  for,  Rana  fishen.  Then  across  the  road  we  found  a  large  male  near 
the  spring  hole,  and  a  little  way  down  the  stream  another  small  one  of  the 
same  species.  We  found  tadpoles  in  the  spring  hole.  Two  hundred  and  fifty 
extra  miles  of  desert  tor  a  frog  seem  worth  it  when  you  find  your  game.  Fol- 
lowed one  liny  stream  back  into  the  field  and  returned  with  the  trophies,  6 
R.  fishen  9  4  Hyla  regilln,  i  Bufo  compactihs.  These  springs  look  marly  or 
alkali.  The  second  frog  we  caught  was  in  one  of  the  headwater  springs  and 
was  resting  against  the  bank  with  i1-:  inches  of  head  out.  When  I  started  to 
catch  it,  it  swam  only  a  short  distance,  didn't  go  into  the  bottom  at  all,  just 
rested  on  top  of  the  bottom.  I  moved  a\va\  the  scum  over  it  and  yet  it  stayed 
in  position.  Then  with  a  pan  we  began  to  scoop  the  area  along  the  edges.  It 


FROGS:  RANIDAE 


455 


came  to  the  top.  They  are  very  easy  to  capture,  not  what  I  would  call  extremely 
alert. 

The  last  one  was  in  the  west  spring  hole.  He  was  deep  in  the  center.  The 
spring  comes  out  here  from  a  deep  hole  under  a  boulder.  These  males  be- 
neath the  surface  or  resting  with  their  heads  out,  when  they  have  no  spots, 
look  much  like  small  R.  catesheiami  but  deeper  green.  After  we  had  caught 
the  first  three,  I  went  up  a  stronger  branch  and  began  to  get  sedgy  and  bushy 
areas.  There  1  found  several  Rana  fishcn  and  Hvla  regtllu.  This  broad  spring 
area,  sedgy  and  shady,  has  been  considerably  cow  punched.  A  little  farther 
on  in  a  heavy  sedge  clump  beside  flowing  water,  two  frogs  leaped  out  at 
once.  I  had  to  catch  one  with  the  right  and  one  with  the  left  hand  at  the  same 
time.  The  water  was  very  cool  in  these  springs. 

Size:  Adults,  i:h-3  inches.  Males,  44-64  mm.  Females,  46-74  mm. 

J.  R.  Slevm  gives  measurements  for  six  adult  specimens  as  45,  48,  51,  56,  57, 
and  60  mm. 

General  appearance:  This  spotted  frog  with  light  stripes  along  the  dorso- 
lateral  folds  is  very  similar  to  R.  pipicn f,  R.  ptdttstns,  or  /?.  scptentrtonahs. 
The  chamois  or  honey  yellow  of  the  hind  limbs  is  very  prominent  in  both 
male  and  female,  reminding  one  of  R.  pulustns.  The  females  are  more  spotted 
than  the  males.  The  upper  parts  arc  olive  green,  the  males  showing  a  tendency 
toward  uniform  color  in  the  forward  part  of  body.  This  forward  part  may 
be  a  brighter  green.  In  the  groin  and  on  the  front  and  rear  of  the  hind  limbs 
are  many  reticulations  of  deep  olive  and  pale  olive-gray. 

Color:  Las  Vegas,  Ncv.  Fcmitle.  Upper  parts  dusky  olive  green,  spots  dark 
greenish  olive,  costal  fold  yellowish  olive.  Sides  are  light  grayish  olive  with 
dark  olive  spots.  In  the  groin  and  on  front  of  hind  legs,  and  rear  as  well,  are 
vcrmiculations  of  deep  olive  and  pale  olive-gray.  Throat  light  grape  green 
or  light  turtle  green  with  some  pale  pinkish  cinnamon,  clouded  with  dark 
grayish  olive.  Breast  and  belly  pinkish  cinnamon  clouded  like  throat.  The 
rear  of  tibia  and  some  of  foot  with  deep  colonial  bufT  or  colonial  buff.  Another 
female  has  throat  cartridge  bufT,  and  breast  and  forelegs  the  same  with  no 
clouding.  Females  have  more  spots  on  the  back  than  the  males. 

Male.  They  range  in  color  from  cedar  green  to  dark  greenish  olive.  The 
spots  range  from  ramette  green  to  pois  green.  The  greatest  difference  between 
males  and  females  is  this  tendency  of  old  males  to  have  spots  obscured,  being 
almost  uniform  like  a  bullfrog.  The  forward  part  of  the  body  may  be  methyl 
green.  The  tympanum  may  be  like  background  or  may  be  wood  brown. 
Honey  yellow  or  chamois  color  is  present  on  underside  of  hind  legs. 

"Color  above  is  brown,  gray,  olive,  or  green,  with  large  or  small  discrete, 
dark  brown  spots  on  head,  body  and  limbs.  These  spots  usually  arc  indefinitely 
bordered  with  light  blue,  gray,  yellow,  or  green,  and  are  irregularly  rounded. 
They  may  form  longitudinal  rows,  or  the  spots  on  their  light  borders  may  be 
nearly  absent.  The  dorso-lateral  fold  may  be  light  or  dark  as  the  general 


456  HANDBOOK  OF  FROGS  AND  TOADS 

ground  color.  Posterior  surface  of  thigh  may  be  more  or  less  clouded,  spotted 
or  marbled  with  brown  or  gray.  Lower  surfaces  white  or  yellow,  sometimes 
clouded,  marbled  or  reticulated  with  gray  or  brown,  especially  on  the  throat" 
(J.  R.  Slevin,  B.C.,  1928,  pp.  126-127). 

Structure:  Thumb  enlarged  in  males;  tympanic  disk  large,  and  larger  in 
males  than  females.  Old  males  may  be  smooth,  but  younger  ones  may  be  more 
warty  than  corresponding  females.  Fourth  toe  with  two  digits  free. 

Dr.  Stejneger  (Calif.,  1893,  p.  227)  gives  the  following  description:  "Heel 
of  extended  hind  limb  reaching  anterior  eye  canthus,  falling  considerably 
short  of  tip  of  snout;  vomerine  teeth  between  and  projecting  posteriorly  be- 
yond choanae;  no  black  ear  patch;  vertical  diameter  of  tympanic  disk  greater 
than  distance  between  nostrils  and  eye;  hind  feet  webbed  for  about  two-thirds; 
one  small  metatarsal  tubercle;  one  weak  dorsolateral  dermal  fold,  no  dorsal 
folds  between;  posterior  lower  aspect  of  femur  granular;  back  and  sides  with 
numerous  small,  distinct,  dark  spots,  surrounded  by  lighter;  no  external  vocal 
sacs.  Habitat:  Vegas  Valley,  Nevada.  Type — U.S.  Nat.  Mus.  No.  18957; 
Vegas  Valley,  Nevada,  March  13,  1891;  V.  Bailey  coll." 

"Head  as  broad  as,  or  broader  than,  long,  slightly  depressed;  snout  rounded, 
projecting  more  or  less  beyond  mouth;  canthus  rostralis  indistinct;  loreal 
region  concave;  nostril  nearer  to  eye  than  to  tip  of  snout;  distance  between 
nostrils  greater  than  mterorbital  width.  Interorbital  width  less  than  width 
of  upper  eyelid.  Tympanum  large,  distinct,  nearly  smooth,  one-half  to  once 
diameter  of  eye.  Fore  limbs  heavy,  no  rudiment  of  pollex;  digits  rather  long, 
first  as  long  as,  or  a  little  longer  than,  second,  third  much  the  longest;  no 
slight  fold  along  sides  of  fingers;  subarticular  tubercles  small  or  moderately 
large,  rounded,  single.  No  tarsal  fold;  inner  metatarsal  tubercle  elongate, 
fairly  prominent;  no  outer  metatarsal  tubercle;  digits  moderately  long;  web 
moderately  full,  one  or  two  phalanges  of  fourth  toe  free.  Skin  above  smooth, 
or  with  a  few  tubercles  or  ridges  on  back  and  sides  of  body  and  dorsal  surface 
of  hind  limbs;  a  strong  dorso-latcral  fold;  a  fold  or  series  of  warts  from  upper 
hp  to  side  of  neck  or  shoulder.  Vomerine  teeth  in  small,  rounded  groups  or 
short  oblique  series  between  or  a  little  behind  the  choanae"  (J.  R.  Slevin,  B.C., 
1928,  p.  126). 

Voice:  These  frogs  have  vocal  sacs  like  Rana  piptens. 

Aug.  20,  1925,  Las  Vegas,  Nev.  When  I  picked  up  no.  10,  he  began  tq  swell 
out  his  sacs  back  of  each  ear  like  the  sacs  of  1?.  pipiens  and  R.  virgatipes.  I  had 
put  the  frogs  in  a  bag  and  carried  them  on  my  belt.  A  few  times  they  gave 
scmicroaks  which  reminded  me  in  some  ways  of  R.  palustris.  Once  or  twice 
heard  a  very  low  croak.  It  must  have  been  Rana  fishen.  Occasionally  in  al- 
coholic males  (MCZ  no.  4840)  one  can  clearly  see  between  the  ear  and  the 
shoulder  the  collapsed  sac. 

Breeding:  This  species  probably  breeds  in  the  spring.  The  eggs  are  not 
known.  The  general  color  of  the  tadpole  is  dull  citrine  with  the  tail  pale  green- 


FROGS:  RANIDAE  457 

yellow,  heavily  mottled.  The  tail  is  elongate,  its  tip  rounded,  length  to  85  mm., 
venter  semitransparent.  The  largest  tadpole  we  caught  was  probably  not  ma- 
ture, being  i  %  inches  (42  mm.)  and  of  the  R.  damitans  type.  The  tooth  ridges 
are  %.  We  caught  no  specimens  at  transformation.  The  smallest  frogs  we  saw 
were  i%(5-i%  inches  (39-42  mm.).  J.  M.  Linsdale  secured  transformation 
sizes  of  28.0-30.5  mm.,  and  from  Las  Vegas  (May  i,  1913)  J.  R,  Slevin  has  a 
30-mm.  transformed  specimen. 

Journal  notes:  Feb.  16,  17,  1942,  Berkeley,  Calif.,  MVZ.  Worked  on  Rana 
fisheri. 

May  16.  What  frog  hunters  we  are!  I  thought  I  was  good  at  it.  I  came  here 
once  with  a  golden  spoon  in  my  mouth.  Seventeen  years  have  gone  since  we 
were  here  last.  Las  Vegas  has  grown,  but  how?  Thirty-five  men  sleeping  on 
the  Union  Pacific  lawn.  Roads  are  changed.  Took  us  most  of  the  day  to  locate 
where  the  old  artesian  well  and  the  springs  were.  At  the  U.S.  Fish  Hatchery 
found  bullfrogs.  The  municipal  golf  course  and  possibly  the  hatcheries  are 
where  the  springs  were.  Looked  these  over  but  no  R.  fisheri.  Tried  Las  Vegas 
Creek  upper  stretches.  Found  a  minnow  and  plenty  of  crayfish  but  no  frogs. 

May  17.  Went  out  Main  Ave.  to  U.S.  Fish  Hatchery.  Looked  around  the  big 
pond.  No  frogs.  Walked  from  municipal  golf  course  along  water  to  main 
Tonopah  road.  Heard  one  jump  in  tules,  probably  my  game.  Went  to  Fifth 
St.  crossing  of  Las  Vegas  Creek.  Looked  it  over.  West  of  this  crossing  in  tules 
heard  one  splash  of  frogs.  Never  saw  them.  This  afternoon  at  4:30  went  to 
Main  Street  crossing  and  walked  up  to  old  artesian  well,  a  mile  or  so.  Some 
minnows  in  stream,  lots  of  crayfish — heard  four  splashes  in  tules  but  never 
saw  frogs.  What  a  state!  Men  sleeping  under  trees,  unemployed,  unhoused 
and  some  unclean,  one  group  quarreling. 

Our  R.  fisheri  may  go  with  the  old  springs  gone,  the  creek  a  mess. 

Authorities'  corner: 
L.  Stejneger,  Calif.,  1893,  p.  227. 
J.  R.  Slevin,  B.C.,  1928,  pp.  126-127. 

"For  many  years  this  form  has  been  considered  a  synonym  of  Rana  onca 
which  was  known  from  the  single  type  specimen  obtained  by  Yarrow  in  1872 
m  'Utah/  So  few  specimens  have  been  available  from  southern  Utah  and 
southern  Nevada  that  the  relationships  of  the  frogs  in  that  area  could  not  be 
studied  satisfactorily.  With  the  series  now  available  from  this  region  along 
with  a  study  of  the  physiography  of  the  area,  I  have  been  able  to  come  to  con- 
clusions as  follows:  The  population  of  frogs  in  Vegas  Valley  is  sharply  isolated 
from  the  closely  related  ones  in  the  Colorado  and  Virgin  River  valleys.  Al- 
though other  colonies  of  Rana  are  isolated  in  eastern  and  southern  Nevada, 
they  have  become  less  markedly  differentiated  from  the  common  type  of 
southwestern  Rana  pipiens.  Whether  the  frogs  of  southern  Utah  and  Nevada 
need  separate  recognition  under  the  name  Onca  as  a  species  or  subspecies  can- 
not be  determined  without  a  study  of  the  whole  species  throughout  its  range, 


458  HANDBOOK  OF  FROGS  AND  TOADS 

but  it  seems  plain  to  me  that  the  Vegas  Valley  ones  should  be  recognized  as 
distinct.  Although  entered  here  as  a  species,  this  form  is  obviously  closely  re- 
lated to  Rana  ptptens  and  it  might  be  well  known  as  a  race  of  that  species.  It 
contrasts  most  sharply  with  that  frog  in  its  peculiar  shade  of  brown  color, 
the  reduction  of  dorsal  spots,  especially  on  the  head,  the  enlarged  tympanum 
and  in  the  reduced  hind  legs"  (J.  M.  Linsdale,  Nev.,  1940,  pp.  210-211). 

Remarks:  Because  of  the  precarious  condition  of  the  limited  Rana  fishen 
habitat,  we  present  this  study  of  the  fine  collections  of  this  species  m  the  Cali- 
fornia Academy  of  Science,  Stanford  University,  and  the  Museum  of  Verte- 
brate Zoology. 

The  California  Academy  of  Science  material  was  all  collected  m  a  small 
stream  from  a  flowing  well  about  a  mile  northwest  of  the  town  of  Las  Vegas. 
Among  the  young  frogs  secured  May  i,  1913,  by  J.  R.  Slcvin,  there  is  a  series 
30,  32,  34,  35  mm.,  some  having  the  transforming  apron  of  the  tadpole  tail. 
In  another  group  collected  Aug.  10, 1913,  by  J.  R.  Slevm  are  males,  all  spotted, 
of  44,  45,  46,  47,  48,  48  mm.;  the  44-mm.  male  has  the  heel  reaching  halfway 
between  eye  and  tip  of  snout.  Nineteen  immature  specimens  31-39  mm.  all 
bear  three  to  four  rows  of  spots.  The  females  range  48,  50, 51, 53,  56,  58, 64  mm., 
some  very  spotted  on  throat  and  on  front  of  femur. 

The  Stanford  University  material  coming  from  Tule  Springs,  north  of  Las 
Vegas,  Nev.,  July  15,  1938,  was  collected  by  Alex  Calhoun,  Robert  Miller, 
and  Carl  L.  Hubbs.  There  are  two  tadpoles,  one  83  mm.  long;  tail  tips  prob- 
ably lost;  labial  teeth  %.  The  upper  and  lower  tail  crests  are  heavily  mottled 
as  in  R.  clamjtans,  much  more  heavily  than  in  our  eastern  R.  pipicns,  but  with 
intestines  showing  through  as  they  do  in  the  latter  form,  not  concealed  by 
pigment  as  in  R.  clamitans  tadpoles.  There  is  one  transforming  frog  measur- 
ing 33.5  mm.,  and  three  females  51,  51,  and  73  mm.,  the  last  with  four  rows 
of  short  folds  between  dorsolateral  folds,  and  sides  with  many  minute  ridges. 

In  the  Museum  of  Vertebrate  Zoology  at  the  University  of  California,  there 
are  among  the  transformed  frogs  collected  by  J.  Linsdale  one  28  mm.  long, 
collected  April  20,  1936,  with  a  tail  stub  of  2.5  mm.,  with  two  irregular  rows 
of  spots  on  dorsum  and  two  or  three  rows  below  dorsolateral  folds,  and  with 
posttympanic  folds  and  numerous  tubercles.  The  fold  of  skin  and  light  stripe 
on  jaw  are  very  obscure,  but  two  folds  forward  from  the  vent  between  dorso- 
lateral folds  are  evident.  Another  frog  secured  May  IT,  1934,  measures  50.5 
mm.,  with  stub  of  tail  at  least  5%  mm.  long  and  three  rows  of  scattered 
spots.  One  of  April  19,  1936,  is  31.5  mm.  and  another  of  May  n,  1934,  is  also 
31.5  mm.  Thomas  Rodgers,  R.  Miller,  and  A.  Calhoun  found  a  very  dark- 
colored  one  36  mm.,  July  15, 1958,  in  which  the  tympanum  is  slightly  inclined, 
two  folds  forward  from  vent  are  made  up  of  tubercles,  and  the  intermediate 
space  ahead  of  these  has  four  irregular  rows  of  these  tubercles  appearing  as 
small  dots.  A  prominent  thin  white  stripe  from  below  the  eye  extends  under 
the  tympanum  and  onto  the  fold  of  skin  back  of  the  tympanum  and  is  fol- 


FROGS:  RANIDAE  459 

lowed  by  an  interrupted  white  mound  at  insertion  of  the  arm.  J.  Linsdale 
collected  one  ^8.5  mm.  April  19,  1936,  and  a  seiies  40,  44,  45.5,  47,  49,  and 
52  mm.  May  i,  1934.  The  44-mm.  frog  has  prominent  light  ridges  on  tarsus 
with  dark  intervals;  the  49-mm.  one  shows  folds  of  the  vocal  sac  but  not  in 
plaits.  It  has  an  enlarged  and  padhke  thumb,  a  slight  suggestion  of  two  rump 
folds,  three  rows  of  spots  between  the  dorsolateral  folds  and  three  below,  but 
all  small.  The  52-mm.  specimen  is  a  male  with  light-colored  vocal  sacs  between 
tympanum  and  brachium,  which  have  rather  deep  valleys  when  not  inflated. 
In  the  collection  is  a  bright  green  female  50.5  mm.  found  by  C.  L.  Camp 
March  2?,  1923.  J.  Linsdale  secured  a  series  of  larger  adult  females,  April  24, 
1936 — 62,  65,  68,  68.5,  and  71  mm.  The  68-mm.  frog  had  two  folds  forward 
from  vent  for  about  %  of  an  inch;  tympana  obliquely  inclined  inward  with  a 
prominent  curved  fold  from  tympanum  to  arm  insertion;  and  two  rows  of 
widely  separated  spots,  five  in  each  row  being  below  the  fold.  In  the  7i-mm. 
female  the  tympanum  was  quite  oblique  mesad  and  a  fold  from  the  inner 
hand  tubercle  to  tibia  was  present  with  a  similar  somewhat  flattened  fold 
from  base  of  fifth  toe  to  the  tibia;  numerous  pebbly  eminences,  some  longi- 
tudinal, are  evident  between  dorsolateral  folds;  and  fold  from  lower  edge  of 
tympanum  toward  shoulder  is  very  prominent  and  unbroken.  The  very 
pebbled  sides  below  dorsolateral  folds  are  grayish  with  two  rows  of  spots 
forward  and  spotting  on  sides  scanty;  reticulations  in  inguinal  region  and 
front  of  femur  are  prominent;  a  spot  is  present  at  insertion  of  arm  and  ex- 
tending along  brachium  to  beginning  of  the  forearm.  The  last  two  joints  of 
fourth  toe  are  free  of  webbing. 

Southern  Bullfrog,  Joe  Brown  Frog,  Pig  Frog,  Swamp  Bullfrog,  Bonnet's 
Frog,  Lake  Frog,  Green  Bullfrog,  Florida  Bullfrog 

Rana  gryho  Stejnegcr.  Plate  XCVII;  Map  31. 

Range:  Louisiana  to  Florida  and  southern  Georgia. 

Habitat:  Aquatic.  Open  water-lily  prairies,  open  ponds,  or  along  deep- 
wooded,  overflowed  banks  of  southern  nvcis,  among  brush  or  similar  debris, 
or  among  aquatic  vegetation,  like  pickerel  weed,  and  more  especially  near 
bushy  edges.  Exceedingly  shy,  hard  to  catch  except  at  night  with  a  light. 

"Lakes  and  marshes;  common.  Our  most  aquatic  frog"  (O.  C.  Van  Hyning, 
Fla.,  1933,  p.  4). 

See  F.  Harper,  Ga.,  1935,  p.  302. 

"Bonnet  and  water-lily  pads  and  prairies;  in  emergent  vegetation  (Panicttm 
hermitomon,  Pontedcria  cordata,  Sagittana  lanajoha)  along  lake  margins; 
among  water-hyacinths  in  lakes  and  streams;  cypress  swamps.  .  .  .  Almost 
wholly  aquatic;  nocturnal,  but  occasionally  fairly  active  on  dark  days.  Various 
observers  have  commented  on  the  extreme  shyness  of  this  frog  .  .  .  ,  but  my 
observations  do  not  confirm  these  impressions.  On  land  it  is  unwary  to  the 


460 


HANDBOOK  OF  FROGS  AND  TOADS 


point  of  stupidity,  while  in  the  water  I  have  caught  as  many  as  fifteen  or 
twenty  consecutively  without  allowing  one  to  escape.  The  fact  that  it  must  be 
hunted  in  most  cases  in  water  two  to  four  feet  deep  has  no  doubt  contributed 
to  its  reputation  for  wildness.  It  is  true  that  if  regularly  annoyed  by  intruders 


Map  ji 

grylio  soon  learns  to  avoid  capture.  .  .  .  R.  grylto  is  more  buoyant  in  the  water 
than  catesbeiana;  it  usually  floats  with  its  head  and  shoulders  well  above  the 
surface.  I  think  that  the  difference  in  the  degree  of  submergence  of  the 
tympanum  may  account  for  the  fact  that  catesbeiana  is  much  more  difficult 
to  approach"  (A.  F.  Carr,  Jr.,  Fla.,  1940^  pp.  66-67). 
Size:  Adults,  31/4-6%  inches.  Males,  82-152  mm.  Females,  85-161  mm. 


FROGS:  RANIDAE  461 

General  appearance:  This  is  a  large  frog  similar  to  the  common  bullfrog 
and  varies  from  brownish  olive  to  blackish  brown  above,  with  some  promi- 
nent, scattered  black  spots.  The  under  parts  often  have  a  network  of  black, 
brown,  and  yellow,  presenting  a  striking  heavily  mottled  appearance.  It  has 
no  dorsolateral  folds. 

Color:  Okefinokee  Swamp,  Ga.  Male.  April  25,  1921.  Upper  parts  rich 
brown  with  black  spots  obscured.  Tympanum  and  outer  circle  deep  brown, 
center  green  or  dark  green.  Top  of  head  dark  green.  Belly  creamy.  Throat 
lemon  yellow  with  dusky  fine  blotches  on  underside  of  fore  and  hind  limbs. 
Lower  belly  blotched  or  reticulated  with  cream  and  brown.  On  sides  of  body 
and  legs  cream  replaced  by  yellow.  First  finger  of  male  much  enlarged.  In 
another  male  head  deep  bottle  green.  Tympanum  center  black,  middle  ring 
light  brown,  outer  ring  brownish  black.  Throat  almost  orange  yellow  on 
either  side.  On  either  side  of  breast  between  arm  insertions  a  deep  green  area. 
On  rear  of  femur  the  yellow  extends  as  a  yellow  line  with  brown  black  on 
either  side  of  it;  this  line  golden  or  orange  yellow,  more  intense  than  yellowish 
or  cream  of  undersides  of  legs.  In  some  the  black  spots  of  the  back  are  farther 
apart.  Black  spots  of  back  surrounded  by  green  with  centers  bronzy  brown. 
Top  of  toes  brown  and  yellow.  Underside  ot  toes  brown.  In  half-grown  male 
under  color  cream,  slight  wash  of  yellow.  Femoral  stripe  cream,  not  yellow. 
Eardrum  small,  all  brown.  Two  pectoral  patches  dusky,  not  green.  (Non- 
Ridgway). 

Female.  June  21,  1921.  Upper  lip  from  tympanum  to  snout  medal  bronze, 
also  top  of  head;  upper  parts  olive-citrine  with  a  few  dark  spots  on  the  pos- 
terior back  and  groin.  Upper  part  of  legs  Saccardo's  olive  with  cross  blotches. 
Throat  baryta  yellow  or,  better,  straw  yellow.  Where  dorsal  color  reaches 
light  venter,  also  on  fore  and  hind  legs,  there  is  some  clear  dull  green-yellow 
or  light  dull  green-yellow.  Venter  marguerite  yellow,  also  same  color  on  top 
of  toes  and  fingers.  Two  patches  on  either  side  of  throat  and  near  pectoral 
region,  so  prominent  in  the  male,  are  light  yellowish  olive,  not  pronounced 
as  in  male.  No  reticulations  on  venter  except  for  faint  ones  on  underside  of 
hind  legs  and  the  slightest  indications  on  the  groin  region.  Rear  of  femur 
with  one  lone  band  of  seafoam  yellow  with  line  of  quaker  drab  below  and 
dark  quaker  drab  or  black  above.  Iris  with  bright  green-yellow  ring  around 
pupil.  Iris  black  with  lots  of  bright  green-yellow  spots.  Green-yellow  outer 
circle  around  the  black. 

Structure:  Large  frog;  no  dorsolateral  folds;  male  with  internal  vocal  sacs, 
the  inflation  forming  a  flattened  pouch  on  throat  with  extra  inflation  on  either 
side,  giving  the  pouch  a  three-parted  appearance;  more  pointed  head  and 
narrower  snout  than  in  /?.  catesbeiana,  and  with  all  the  hind  toes  except  the 
fourth  proportionally  longer;  webs  to  the  tips  of  toes. 

"The  most  obvious  difference  between  this  species  and  the  ordinary  bull- 
frog (Rana  catcsbeiana)  is  the  great  length  of  the  toes,  except  the  fourth,  the 


462  HANDBOOK  OF  FROGS  AND  TOADS 

latter  consequently  projecting  much  less  beyond  the  others  than  in  R.  cates- 
betana,  in  which  the  third  toe,  measured  from  the  inner  metatarsal  tubercle, 
is  considerably  less  than  three  and  one-half  times  the  difference  between  it  and 
the  fourth  toe.  In  fact,  this  difference  is  seldom  more  than  one-fourth  m  the 
new  species  and  seldom  less  than  one-third  in  R.  catesbetana. 

"In  order  to  ascertain  exactly  the  proportions  of  the  first  four  toes  in  both 
species  large  series  of  both  species  were  measured,  viz.:  12  of  Rana  grylio 
and  50  of  R.  catesbeiana,  the  measurements,  as  well  as  their  equivalents  ex- 
pressed in  percentages  of  the  fourth  toe,  being  given  at  the  end  of  this  article. 
In  order  to  get  as  stable  a  starting  point  as  possible  for  these  measurements 
the  anterior  edge  of  the  inner  metatarsal  tubercle  was  chosen  and  the  length 
of  the  toe  in  this  case  consequently  means  the  distance  from  this  point  to  the 
tip  of  the  toe  in  question. 

"The  proportions  obtained  in  this  way  may  be  expressed  as  follows: 

Rana  grylio  Rana  catesbeiana 

Per  cent  of  Per  cent  of  fourth 

fourth  toe  toe 

Third  toe                      80  to  84  70  to  76 

Second  toe                     55  to  61  47  to  51 

First  toe                        34  to  39  27  to  33 

There  is  consequently  no  overlapping  or  intergrading.  The  fourth  toe  has 
the  same  relative  length  in  both  species,  but  in  the  new  one  the  other  toes 
have  become  considerably  lengthened,  thus  giving  a  much  larger  surface  of 
web  than  in  the  ordinary  bullfrog.  .  .  . 

"The  range  of  the  new  species  is  as  yet  known  but  fragmentarily,  as  we  have 
specimens  only  from  southern  Florida,  Pensacola,  Florida,  and  Bay  St.  Louis, 
Mississippi.  The  habitat  of  R.  grylto  is  thus  partly  occupied  by  R.  catesbeiana, 
which  certainly  occurs  in  northern  Florida  and  on  the  Gulf  coast  reaches  New 
Orleans.  The  overlapping  of  the  two  forms  affords  additional  evidence  of 
their  specific  distinctness,  if  such  were  needed.  It  may  be  mentioned  in  this 
connection  that  the  most  southern  specimens  of  R.  catesbetana  do  not  show 
the  slightest  tendency  of  a  variation  toward  R.  grylio,  as  is  clearly  proven  by 
the  measurements  .  .  ."  (L.  Stejneger,  Fla.,  1901,  pp.  212-215). 

Voice:  The  call  is  a  grunt  like  that  of  a  pig  or  an  alligator.  It  is  rough  and 
guttural. 

Dr.  E.  A.  Mearns  says  it  resembles  "the  grunt  of  a  herd  of  pigs."  To  Dr. 
Harbour  in  1920,  "its  call,  which  is  heard  at  night,  or  on  damp  days,  resembles 
the  grunting  of  a  pig,  and  consists  of  but  one  sound  oft  repeated."  Dr.  J.  C. 
Bradley  called  its  note  "peculiar  alligatorlike  grunts." 

In  our  journal  under  June  18, 1912,  we  wrote:  We  heard  plenty  of  southern 
bullfrogs  from  8-n  A.M.  The  water  was  waist  deep  and  the  bushes  8-12  feet 
above  the  water.  One  of  their  notes  was  very  much  like  that  of  Rana  cates- 


FROGS:  RANIDAE  463 

beiana  of  the  North.  Another  note  frequently  given  in  chorus  was  mournful, 
reverberating,  prolonged,  and  deep,  not  at  all  like  the  regular  bullfrog  chorus 
note.  There  were  a  number  in  the  chorus  and  it  sounded  like  one  continuous 
long  deep  roar.  The  following  day,  June  19,  from  7-11  A.M.,  when  the  sun 
was  very  hot,  we  heard  several  along  the  tree-covered  overflowed  banks  of 
Suwannee  River.  On  June  13,  while  traveling  through  Minne  Lake  Run  we 
heard  the  deep  guttural  notes  of  the  "Swamp  Bullfrog"  all  along  the  bonnet 
borders  and  through  the  cypress  woods  on  either  side. 

The  male  frog  has  internal  vocal  sacs  like  Rana  clamitans  and  Rana  cates- 
betana,  but  the  inflation  is  greater  than  in  these  two  species.  There  is  a  central 
inflation,  but  on  either  side  of  the  middle  is  an  extra  inflation,  giving  the  sac 
a  three-parted  appearance.  The  croak  can  be  heard  at  considerable  distance 
and  a  midnight  chorus  is  loud.  They  call  spread  out  on  the  surface  of  the 
water. 

June  12,  1917,  two  miles  from  Theodore,  Ala.,  we  heard  the  croak  or  grunt 
(4  or  5  notes)  of  the  southern  bullfrog:  hard  to  find,  lost  the  first  one.  They  are 
among  cat  briars  (Smthuc).  Found  a  young  one  and  later  took  a  large  male. 
To  one  member  of  the  paity  there  seemed  something  human  in  its  voice. 

Breeding:  They  breed  from  March  r  through  September,  generally  in 
humid  weather,  with  night  temperatures  of  (^°-jo°  and  day  temperatures  of 
85°-9o°.  The  crest  is  in  June  and  July.  The  black  and  while  eggs  arc  in  a  sur- 
face film  12  by  12  inches  to  12  by  25  inches,  and  attached  to  vegetation.  The 
egg  is  y\\\  inch  (1.6  mm.),  the  outer  envelope  is  \{\-\\  inch  (4-6  mm.),  merg- 
ing into  the  jelly  mass.  The  greenish  tadpole  is  quite  large,  4  inches  (100  mm.), 
with  tail  long  and  sharply  acuminate.  The  tooth  ridges  are  %.  After  a  tadpole 
period  of  i  to  2  years,  the  tadpoles  transform  fiom  April  24  to  July  19,  at 
i%-2  inches  (52-49  mm.).  A  young  transformed  R.  grylio  looks  much  like 
an  adult  or  half-grown  carpenter  frog  (R.  virgatipes). 

"They  call  every  month  of  the  year"  (A.  F.  Carr,  Jr.,  Fla.,  i^ob,  p.  67). 

Journal  notes:  1912,  Okcfinokee  Swamp,  CJa.  I  will  never  forget  my  first 
capture  of  the  swamp  bullfrog.  The  prominent  black  or  brown  and  yellow 
reticulations  of  the  under  parts  are  very  conspicuous.  Sometimes  we  succeeded 
in  killing  them  with  an  oar  or  pole  while  pushing  through  the  bonnets  and 
occasionally  the  members  of  the  party  captured  them  when  visiting  set  trap 
lanterns  for  insects. 

On  April  25,  1921,  we  went  with  flashlight  after  Rana  grylio.  On  the  vegeta- 
tive carpet  and  lily  pads  were  untold  numbers  of  Acns,  on  the  lily  pads  and 
on  the  bushes  Hyla  ctnerea,  and  among  the  pickerel  weeds  different  sizes  of 
Rana  grylio.  In  all,  three  of  us  secured  only  six  adults  and  one  transformed 
specimen.  Later  I  found  that  they  could  be  picked  up  rather  easily  with  a 
flashlight.  On  May  6,  Dave  Lee  went  out  on  Billy's  Lake  at  night  with  a  torch 
to  catch  small  fish  for  bait.  With  a  small  clip  net  he  would  scoop  in  front 
of  a  Rana  grylio  and  usually  catch  it. 


464  HANDBOOK  OF  FROGS  AND  TOADS 

On  May  10, 1921,  we  found  one  cypress  pond  with  plenty  of  Rana  grylio  in 
all  stages.  Our  comment  is  that  they  are  easy  of  capture  at  night  except  for 
adults  and  even  these  can  be  taken.  On  May  n  we  found  many  R.  grylio  at 
night  along  the  edges  of  Billy's  Lake.  They  usually  were  at  the  edge  near  the 
bushes  or  under  them  or  among  brush  or  in  maiden  cane.  When  one  tried 
to  take  their  pictures,  they  were  too  much  surrounded  with  vegetation  or 
sticks.  We  could  approach  closely,  but  in  clearing  away  brush  for  the  flash- 
light picture  we  would  scare  them.  Often  we  heard  them  go  skipping  along 
across  the  lake  or  along  the  edges  like  Rana  catesbeiana.  Like  most  shy  frogs 
they  can  be  captured  if  females  are  around.  In  some  cases  the  females  led  the 
way  to  where  a  male  might  be.  Where  we  succeeded  in  flashlighting  a  croak- 
ing male,  we  were  quite  certain  females  were  near  because  of  eggs  found 
there  soon  afterwards.  On  June  20  we  found  another  method  of  capture.  We 
never  dreamed  it  would  work  for  Rana  grylto.  We  would  pull  in  hurriedly 
masses  of  floating  maiden  cane,  and  in  the  corner  at  the  bottom  of  the  boat 
all  kinds  of  life  would  drop.  On  this  night  we  caught  two  Rana  grylio  speci- 
mens, one  a  fine  ripe  female.  All  in  all,  our  main  reliance  was  to  wade  about 
at  night  on  the  prairies  or  in  open  cypress  ponds  and  to  catch  them  by  hand. 

"Anent  this  species  it  might  be  interesting  to  include  some  comments  of 
Percy  Viosca,  Jr.,  in  a  letter  to  Dr.  H.  F.  Moore,  U.S.  Bureau  of  Fisheries. 
On  July  ii,  1923,  he  writes:  *It  might  also  interest  you  to  know  that  Rana 
grylio  is  abundant  in  this  State  [  La.|,  but  we  have  found  it  is  a  thoroughly 
aquatic  species  and  not  very  hardy  in  captivity,  and  would  not  be  suitable 
for  frog  culture  even  though  it  is  a  fine  flavored  species  and  we  believe  much 
superior  to  flesh  of  the  catesbeiana.  It  would  be  only  suitable  for  planting  in 
shallow  lakes  of  the  lagoon  type,  but  once  established  it  would  be  difficult  to 
collect,  except  by  experienced  night  hunters.  It  would  be  a  species  very  dif- 
ficult to  ship.  Although  we  have  shipped  them  successfully,  the  frogs  need 
individual  care*  "  (A.  H.  Wright,  Gen.,  1932,  pp.  $69-370). 

Authorities'  corner: 
L.    Stejneger,    Fla.,    1901,    pp.    212-      P.  Viosca,  Jr.,  La.,  1918,  p.  161. 

213.  P.  Viosca,  Jr.,  La.,  19233,  p.  39. 

R.  F.  Deckert,  Fla.,  1914,"*  pp.  3-4.  A.  H.  Wright,  Gen.,  1932,  pp.  368-369. 

M.  C.  Dickerson,  Gen.,  1914,  pp.  226- 

227- 

"The  largest  frog  known  in  Florida  and  on  the  sea  coast  of  Carolina  f  fhainly 
R.  grylio  but  the  Carolina  ones  may  be  also  R.  catesbeiana  \  is  about  eight  or 
nine  inches  in  length  from  the  nose  to  the  extremity  of  the  toes;  they  arc  of 
a  dusky  brown  or  black  colour  on  the  upper  side,  and  their  belly  or  under 
side  white,  spotted  and  clouded  with  dusky  spots  of  various  size  and  figure; 
their  legs  and  thighs  also  are  variegated  with  transverse  ringlets,  of  dark 
brown  or  black,  and  are  yellow  and  green  about  their  mouth  and  lips;  they 
live  in  wet  swamps  and  marshes,  on  the  shores  of  large  rivers  and  lakes;  their 


FROGS:  RANIDAE  465 

voice  is  loud  and  hideous,  greatly  resembling  the  grunting  of  a  swine,  but 
not  near  as  loud  as  the  voice  of  the  bullfrog  of  Virginia  and  Pennsylvania, 
neither  do  they  arrive  to  half  their  size,  the  bull  frog  being  frequently  eighteen 
inches  in  length,  and  their  roaring  as  loud  as  that  of  a  bull"  (W.  Bartram, 
Gen.,  1791,  pp.  276-277). 

"It  has  an  enormous  head,  a  thick-set  body,  and  strongly  formed  limbs, 
much  thicker  in  proportion  than  those  of  the  common  European  frog.  Its 
croak  is  spoken  of  as  'booming1  and  is  very  loud.  It  can  sometimes  be  heard 
appearing  to  come  from  every  part  of  an  extensive  district,  and  from  a  mile 
or  more  distant.  I  am  not  sure  that  (he  Okefinoke  frog  is  not  a  distinct  variety 
of  the  common  bull-frog.  It  is  larger  than  those  which  I  have  seen  in  streams 
running  into  the  Mississippi  and  its  tributaries,  and  in  other  places,  and  seems 
to  differ  in  colour,  though  neither  of  these  circumstances  is  of  much  im- 
portance. Size  may  depend  on  locality  and  abundance  of  food,  and  coloration 
certainly  does.  The  following  are  the  measurements  of  the  largest  specimen 
that  I  could  find.  Length  of  body  from  nose  to  base,  S  inches;  breadth  across, 
nearly  5  inches;  girth,  14  inches;  length  ot  hind-limb,  i2!/2  inches;  weight, 
within  a  fraction  of  three  pounds.  The  usual  size  is  about  seven  inches  in 
length,  half  as  much  more  for  length  of  hind-limb,  and  a  weight  of  over  two 
pounds"  (P.  Fountain,  Ga.,  1901,  pp.  62-63). 

River-Swamp  Frog,  Greenback,  Heckscher's  Frog,  Wright's  Bullfrog, 

Alligator  Frog 

Rana  hecfocAen  Wright.  Plate  XCVIII;  Map  32. 

Range:  Coastal  South  Carolina  to  Oklawaha  River,  Fla.,  west  to  Biloxi, 
Miss.,  and  possibly  to  Louisiana  and  North  Carolina. 

"To  date  this  Lower  Austral  species,  smaller  than  the  common  bullfrog, 
has  been  recorded  as  far  north  along  the  coast  as  Charleston  County.  W.  W. 
Humphreys  took  several  specimens  from  a  flooded  sand  pit  near  the  Edisto 
River  at  Pon  Pon,  Charleston  County,  June  1935"  (E.  13.  Chamberlain,  1957). 

"In  July  1958,  John  C.  Pearson  presented  the  Charleston  Museum  with  a 
number  of  strikingly  marked  tadpoles  taken  at  Hail's  Bluff,  Edisto  River, 
Dorchester  County.  Ivan  R.  Tomkins  sent  in  two  specimens  from  Canston's 
Bluff  near  Savannah,  Georgia,  in  February  1935"  (E.  B.  Chamberlain,  S.C., 

19 $9,  PP-  30-3')- 

Habitat:  Swampy  edges  of  rivers  and  streams,  a  truly  fluviatile  species. 

"I  once  took  twenty  adults  together  with  eighteen  catcsbeiana  in  a  swamp 
near  the  Santa  Fe  River  (Columbia  County).  .  .  .  Nocturnal,  terrestrial. 
Where  hec^schen  and  cate*beiana  occupy  the  same  major  habitat  they  are 
found  in  essentially  the  same  minor  habitats,  and  apparently  have  very  similar 
habits.  Three  or  four  individuals  of  each  species  are  often  found  seated  about 
residual  pools  along  the  course  of  an  intermittent  swamp  stream  near  Gaines- 


466 


HANDBOOK  OF  FROGS  AND  TOADS 


Plate  XCVII.  Rana  grylio.  i.  Female 
(XH).  2.  Male  croaking  (X%).  3,4-  Fe- 
males 


XCF///.  fla«a  hec^scheri.  1,4.  Fe- 
males (X%).  2.  Tadpole  (XVi).  3.  Male 
(X%).  5-  Hind  foot  (X%).  6.  Young  tad- 
pole (x%). 


FROGS:  RANIDAE 


467 

ville.  The  two  show  marked  difference  in  their  reactions  to  the  presence  of 
an  intruder,  hecfahen  being  much  more  phlegmatic  in  temperament;  it  is 
not  alarmed  easily  and  shows  little  shrewdness  in  eluding  capture.  When 


Map  32 

caught,  it  hangs  limp  in  the  hand,  never  resorting  to  the  vigorous  kicks  and 
contortions  characteristic  of  grylio  and  catesbeiana"  (A.  F.  Carr,  Jr.,  Fla., 
i94ob,  p.  66). 

Size:  Adults,  3!4~5  inches.  Males,  82-151  mm.  Females,  102-150  mm. 

General  appearance:  This  is  a  large  greenish  black,  rough-skmncd  frog 
with  the  venter  very  heavily  mottled  with  black.  The  throat  of  the  male  has 


468  HANDBOOK  OF  FROGS  AND  TOADS 

a  wash  of  yellow  on  a  light  gray  background.  The  edge  of  the  lower  jaw  with 
light  spots  of  white,  yellow,  or  buff,  is  similar  to  that  of  the  green  frog.  The 
center  of  the  eardrum  is  rough,  light  brown  or  greenish  buff,  with  the  re- 
mainder very  dark,  almost  black.  The  webbing  on  the  foot  does  not  extend 
to  the  tip  of  the  fourth  toe,  leaving  about  i1/^  joints  free.  Rear  of  femur  with 
prominent  light  spots;  its  color  pattern  not  an  alternation  of  light  and  dark 
parallel  bars  as  in  R.  grylio,  R.  virgattpes,  and  Acris  gryllus. 

Color:  Callahan,  Fla.  Adult  male  (95  mm.).  Aug.  18,  1922.  General  dorsal 
color  citrine-drab  to  grayish  olive  becoming  on  top  and  sides  of  head  and 
center  of  tympanum  dark  olive-buff,  isabella  color,  or  cinnamon-brown.  Ear- 
drum except  middle  mummy  brown.  On  back  of  body,  on  head,  and  on  sides 
is  some  serpentine  green.  Under  parts  spotted  white,  with  glaucous  gray, 
light  Payne's  gray,  or  pale  drab  on  throat  and  breast.  Throat  with  a  little 
citron  green  or  deep  chrysolite  green.  Spots  on  lower  jaw  rim  four  or  five, 
seafoam  yellow  to  deep  colonial  buff  in  the  spot  just  back  of  the  angle  of  the 
mouth.  This  spot  except  above  surrounded  by  black.  Black  spot  just  below 
angle  of  mouth  to  and  across  the  insertion  of  the  brachium.  Three  black  spots 
on  the  front  edge  of  the  antebrachium.  Rear  of  forelegs  black  to  tips  of  fingers 
and  webs.  Tops  of  the  fingers  with  seafoam  yellow  or  deep  colonial  buff  spots. 
Narrow  black  bars  across  the  dorsum  of  the  femur,  tibia,  and  hind  foot.  Rear 
of  femur  with  white  unconnected  spots  on  the  bone  brown  ground  color.  Rear 
edge  of  hind  foot  to  tip  of  fourth  and  fifth  toes  black.  Outside  rim  of  ins  bright 
green-yellow,  inner  rim  capucme  orange;  interval  black  with  orange-rufous 
spots. 

A  younger  specimen,  65  mm.  long,  is  dark  olive  or  deep  olive  on  entire 
upper  parts  and  very  warty.  Throat  is  deep  grayish  olive. 

Another  male.  June  10,  1928.  When  first  caught  this  frog  was  very  black, 
and  white  beneath,  with  throat  washed  with  oil  yellow  on  light  mineral  gray 
ground.  The  markings  of  the  femur  are  in  spots  of  white  or  pale  vinaceous- 
fawn  in  contrast  with  the  lines  of  white  conspicuous  m  Rana  grylio. 

Another  male  is  olive  or  dark  greenish  olive  above. 

Adult  female.  June  10,  1928.  Much  the  same  as  male  but  upper  parts  much 
more  spotted  with  black.  Head  spotted.  Tympanum  with  blotches  of  black 
or  olivaceous  black  (3).  Half  bars  on  femur.  Bar  across  arm  insertion.  Throat 
with  no  or  little  oil  yellow  or  sulphme  yellow.  White  spots  or  pale  vmac^ous- 
fawn  spots  on  rim  of  lower  ]aws  prominent.  Under  parts  not  quite  so  spotted. 
Iris  same  as  in  male. 

Young  transformed  frogs.  June  10,  1928.  A  young  transformed  frog  has  a 
prominent  eye.  Sometimes  brick  red.  Often  mars  orange  predominates  over 
black,  and  iris  rim  is  orange-rufous  or  brighter.  Entire  under  parts  finely 
speckled  white  and  deep  olive-gray  or  deep  grayish  olive. 

Structure:  Upper  parts  sometimes  very  warty;  no  lateral  fold;  center  of  ear- 


FROGS:  RANIDAE  469 

drum  granular  and  light  in  color;  males  with  tympanum  much  larger  than 
eye;  thumb  enlarged,  and  throat  darker  and  with  yellow;  back  more  warty 
in  male. 

"Like  Rana  gryho  and  Rana  catesbeiana  it  has  no  dorsolateral  fold  and  no 
phalanx  of  the  fourth  toe  is  totally  free  of  web;  third  toe  in  56  mm.  specimens 
i  to  3  mm.  shorter  in  Rana  hec{schert  than  in  the  other  two  species  of  bull- 
frog, or  3  to  6  mm.  shorter  in  82  mm.  specimens  or  6  to  9  mm.  shorter  in  95 
mm.  specimens;  third  toe  3.8  (95  mm.)-3.56  (82  mm.)-3.3  (56  mm.)  in  length 
(snout  to  vent)  in  R.  hec{scheri  while  2.7  to  3.1  (95-56  mm.)  in  R.  grylio  and 
R.  catesbeiana;  third  toe  1.6-1.7  in  fourth  toe  in  R.  hec^scheri  while  1.2-1.5 
in  fourth  toe  in  the  other  two  species;  first  finger  decidedly  longer  than  sec- 
ond, while  in  the  other  two  species  it  is  usually  shorter  or  sometimes  equal; 
first,  second,  third  and  fifth  toes  shorter  than  corresponding  toes  of  R.  grylio 
and  R.  catesbeiana;  fourth  finger  8.6  (95  mm.)-8.2  (82  mm.)-8.o  (56  mm.) 
in  length  (snout  to  vent)  while  6.0  to  6.3  (95  mm.)-6.8  to  7.4  (82  mm.)-5.6 
to  7.1  (56  mm.)  in  the  other  two  species;  mternasal  space  less  than  upper  eye- 
lid width,  1.07-1.43  m  it  while  .85-1.0  in  R.  grylio  and  R.  catesbeiana;  tympa- 
num m  males  is  proportionally  greatest  in  R.  grylto,  somewhat  smaller  in 
R.  catesbeiana  and  R.  clamitans  and  smallest  in  JR.  hecfocheri;  intertympanic 
width  of  95  mm.  males  in  length  (snout  to  vent)  4.52  in  R.  hec^schen,  5.4$ 
in  R.  clamitans  and  6.3  in  R.  grylio  (R.  catesbeiana  males  of  95  mm.  have 
tympanum  poorly  developed,  but  a  1 36  mm.  male  has  it  4.85) :  in  general, 
intertympanic  width  broadest  in  R.  hecl^schcn  and  R.  catesbeiana  and  nar- 
rowest in  R.  grylio;  distance  from  the  rear  corner  of  the  eye  to  the  same  corner 
of  the  other  eye  much  greater  than  the  intertympanic  width  in  R.  gryltot 
somewhat  greater  in  R.  clanntans,  about  equal  in  R.  catesbeiana,  and  equal  in 
1?.  hecl(schen,  i.e.,  in  the  males. 

"In  spirits,  four  95  mm.  males  of  four  species  are  as  follows:  bister  or 
mummy  brown  on  dorsum  of  R.  catesbeiana,  brownish  olive  in  R.  grylto,  deep 
grayish  olive  in  R.  clamitans,  and  deep  mouse  gray  in  R.  hecl^scheri;  upper 
parts  without  very  distinct  dark  spots  m  R.  clamitans  and  R.  catesbeiana,  with 
prominent  large  black  spots  m  R.  gryho,  and  with  many  small  dark  spots  in 
R.  hecl(scheri;  venter  of  R.  damttant  clear  white  except  for  the  yellow  throat, 
venter  of  R.  catesbeiana  heavily  blotched  with  black,  so  also  m  R.  grylio — all 
three,  however,  with  a  white  background  color  but  in  R.  hecl^scheri,  the  deep 
mouse  gray  or  dark  color  so  prominent  it  becomes  the  background  color  and 
the  white,  scattering  spots;  light  spots  on  upper  and  lower  Jaws  more  promi- 
nent than  in  R.  clamitans"  (A.  H.  Wright,  Ga.,  1924,  pp.  145-144). 

Voice:  One  call  is  a  snore  or  snort,  and  another  a  peculiar,  snarling,  ex- 
plosive grunt.  The  discovery  of  this  frog  came  from  his  voice.  On  Aug.  18, 
1922,  we  visited  Alligator  Swamp  at  Callahan,  Fla.,  because  of  some  curious 
large  tadpoles  with  black  borders  on  their  tails  that  we  had  seen  one  month 


470  HANDBOOK  OF  FROGS  AND  TOADS 

earlier.  Mrs.  Wright  discovered  a  queer-looking  green  frog,  as  she  supposed, 
and  was  calling  to  us  when  we  were  startled  by  a  call  unlike  any  other  Rana 
we  had  ever  heard. 

For  other  notes  see  Harper,  Gen.,  195?,  pp.  302-306. 

"Intermittent  croaks  may  be  heard  during  the  first  warm  rains  in  April 
(Alachua  County),  and  spasmodically  throughout  the  summer"  (A.  F. 
Carr,  Jr.,  Fla.,  1940^  p.  66). 

Breeding:  The  eggs  arc  probably  a  surface  film.  The  tadpole  grows  large, 
3%  inches  (97  mm.).  The  young  tadpole  is  black  with  a  gold  band  across  the 
body  and  the  tail  clear.  The  mature  tadpole  is  dark  in  body  with  black-rimmed 
tail  crests.  The  tooth  ridges  arc  %  or  %.  The  tadpole  period  is  probably  2 
years.  It  transforms  in  June  at  iMj-i%«;  inches  (30-39  mm.),  sometimes  at  2 
inches  (49-52  mm.).  The  small,  transversely  banded  tadpoles  were  very 
numerous  in  June,  1921,  July  17,  1922,  and  June  8-10,  1928.  This  seems  to 
indicate  spring  and  early  summer  as  the  breeding  period — last  week  of  April 
to  July  8. 

Young  tadpole  black  with  a  gold  band  across  the  body  and  tail  clear.  When 
half  grown  this  band  disappears.  The  mature  tadpole  may  become  about  as 
large  (97  mm.)  as  a  bullfrog  tadpole.  The  black-rimmed  crests  of  the  tail  and 
black  bands  on  upper  tail  musculature  and  light  color  cm  lower  half  are  the 
conspicuous  and  distinctive  markings.  Body  dark  greenish  olive  finely  covered 
with  pale  greenish  flecks.  On  the  venter,  vinaceous-fawn,  vmaceous-cmnamon, 
or  orange  vmaceous.  Clump  of  four  to  six  large  dark  spots  on  throat.  Lower 
belly  pale  forget-me-not  blue,  upper  belly  and  breast  jay-blue  to  grayish 
violaceous  blue.  Spots  of  back  thicker  on  lower  belly.  Lateral  line  of  pores 
conspicuous  on  head  and  body. 

We  started  on  this  winding  "tadpole  trail"  on  our  first  trip  to  Okefinokee 
Swamp,  Ga.,  in  1912.  July  17,  1922,  in  a  cut-off  overflow  pool  at  Thompson 
Landing  (south  of  Folkston)  on  St.  Mary's  River  we  found  an  almost  pure 
culture  of  a  small  black  tadpole  with  a  gold  and  white  transverse  band  like 
those  of  June  16,  1912. 

My  journal  reads:  In  one  cut-off  pool  in  a  watercourse  (which  now  is  a 
succession  of  separated  pools)  we  hauled  the  seme.  It  was  covered  with  a 
wriggly  mass  which  at  first  looked  like  water  bettles  to  Miles  (Dr.  M.  D. 
Pirnie).  I  must  confess  I  would  have  seen  them  the  same  way  if  I  had  nctt  had 
experience  with  them  10  years  before.  The  tails  being  transparent  were  hardly 
in  evidence.  They  have  a  band  across  the  back.  In  another  cut-off  from  St. 
Mary's  River  they  were  in  immense  numbers  in  shallow  water  and  presented 
a  very  beautiful  sight  in  the  sunlight  with  their  dark  bodies  and  transverse 
bands.  Were  it  not  for  the  transverse  bands,  they  would  look  like  toad  tad- 
poles. Then  I  provisionally  placed  them  with  the  green  tree  frog,  Hyla  cmerea, 
which  sometimes  has  somewhat  the  same  appearance. 

Three  days  later,  July  20,  we  started  for  Jacksonville,  Fla.  Just  north  of 


FROGS:  RANIDAE  47! 

Callahan,  near  a  large  concrete  bridge,  a  car  was  stuck  on  a  smaller  detour 
bridge  and  we  had  to  wait.  In  the  areas  beside  the  new  Dixie  Highway  were 
shallow  ponds  or  overflowed  areas.  These  were  tributary  to  Alligator  Swamp, 
which  in  turn  is  a  part  of  Mills  Swamp  (U.S.  Gcol.  Survey  Quadrangle— 
Milliard).  At  first  I  saw  a  few  cross-banded  forms  of  the  Thompson  Landing 
sort  that  I  had  taken  to  be  Hyla  cincrca.  I  began  to  suspect  they  might  be 
R.  capita  or  R.  virgatipes,  probably  the  former.  We  collected  a  few  and  went 
on.  The  larger  ones  were  very  conspicuous  with  black-runmed  crests  and 
black  bands  on  upper  half  of  tail  musculature  and  a  light  color  on  lower 
musculature.  In  one-third  grown  tads  the  crossbands  show  through  faintly. 
When  full  grown  they  disappear.  On  July  21,  on  our  return  from  Jackson- 
ville, we  stopped  at  Callahan  (Alligator  Swamp).  The  tadpoles  of  July  20 
were  abundant.  They  travel  in  big  schools  as  no  other  big  tadpoles  do.  They 
remind  me  of  a  school  of  mature  Bitfo  tadpoles.  Once  in  a  while  among  the 
fair-sized  ones  were  monsters  almost  as  big  as  bullfrog  tadpoles.  And  these 
monsters  have  no  suggestion  of  hind  leg  buds.  Does  this  species  winter  over 
i  or  2  years  as  a  tadpole? 

We  checked  up  our  Thompson  Landing  (On.)  and  Alligator  Swamp  (Fla.) 
material  and  found  them  all  of  the  same  species,  and  the  trail  ended  with  the 
naming  of  the  new  frog  Ran  a  hccl(schcn  Wright. 

"The  number  ot  tadpoles  produced  in  a  given  breeding  site  is  astounding. 
Wright  (19^)  remarks  on  their  abundance  at  Callahan.  In  Alachua  County 
they  transform  consistently  in  April  and  May.  H.  K.  Wallace  and  I  found 
them  emerging  in  tremendous  numbers  in  a  backwater  of  the  Santa  Fc  River 
May  i,  ig^;  we  probably  could  have  collected  a  thousand  with  little  difficulty. 
The  disparity  in  relative  abundance  between  the  adults  and  the  young  of  this 
species  indicates  an  extremely  low  survival  potential"  (A.  F.  Carr,  Jr.,  Fla., 
iQ4ob,  p.  66). 

"Rana  hect^schcn  is  very  common  m  the  vicinity  of  Silver  Springs,  Florida, 
in  the  streams  and  rivers  and  in  the  lakes  connected  by  streams  which  flow 
into  the  Oklawaha  River.  The  night  of  June  S,  1936,  I  caught  three  mating 
pairs  and  transferred  them  to  my  private,  twenty  acre,  grassy  lake.  About 
five  thousand  eggs  were  laid  in  masses,  in  shallow  water,  among  the  water 
grasses;  they  were  hatched  in  ten  to  fifteen  days  into  black  tadpoles.  About 
ten  per  cent  of  the  eggs  failed  to  hatch.  From  then  until  September  the  tad- 
poles grew  rapidly,  attaining  the  length  of  four  to  five  inches.  The  color 
changed  to  a  lighter  hue  and  showed  a  speckled  appearance;  the  tail  was 
bordered  with  black.  The  tadpoles  stayed  in  shallow  water  and  were  always 
packed  close  together.  They  fed  on  meat  thrown  into  the  lake  for  the  alli- 
gators, and  seemed  also  to  feed  on  plant  life.  Occasionally  dead  tadpoles  were 
noted  but  I  could  not  determine  the  cause  of  death. 

"The  growth  was  slower  during  the  cooler  months  but  the  tadpoles  re- 
mained very  active.  Banded  water  snakes  fed  on  them  occasionally.  Some- 


472  HANDBOOK  OF  FROGS  AND  TOADS 

times  the  tadpoles  came  into  shallow  water  so  that  their  backs  were  exposed, 
but  at  night  they  went  back  into  deep  water.  They  were  always  found  on 
the  same  side  of  the  lake  within  an  area  of  fifty  yards.  By  March  1937,  the 
hind  legs  appeared  and  growth  was  again  noticeable.  By  April  i  the  tadpoles 
were  six  to  seven  inches  long  and  front  legs  appeared.  On  April  10  the  trans- 
formation was  well  under  way  and  the  tadpoles  were  hopping  out  on  the 
bank.  During  this  time  great  numbers  (about  twenty  per  cent)  died.  By  May 
i  all  had  transformed,  hopped  out  on  the  bank  and  scattered.  On  June  i,  one 
was  found  hopping  away  about  two  hundred  yards  from  the  lake.  Very  few 
can  be  found  now  anywhere  around  the  lake"  (E.  R.  Allen,  Fla.,  1938,  p.  50). 

Journal  notes:  In  August,  1922,  at  Callahan,  Fla.,  we  captured  with  a  light 
eight  or  ten  frogs  of  various  sizes  from  i-year  frogs  to  full-sized  adults.  We 
found  them  in  the  shrubbery  and  on  the  banks  about  the  bases  of  trees.  More 
were  captured  than  lost.  They  were  rather  awkward  in  their  escape  and  would 
tumble  off  their  perches. 

June  8,  1928,  Callahan,  Fla.  This  afternoon  went  to  type  locality  for  Rana 
hect($cheri.  Saw  lots  of  tadpoles  among  vegetation.  Captured  about  12  trans- 
formed ones.  Can  approach  them  when  heads  are  out  of  water  and  can  catch 
them.  Some  rest  on  mud  of  bank.  Many  here  with  four  legs  and  long  tail. 
Went  along  in  pickerel  weed  on  east  side  of  bridge  where  is  shady  at  5  P.M. 
and  finally  saw  a  large  one.  Slowly  approached  from  water  side  with  net  and 
came  close  enough  to  catch  frog.  Started  skirting  several  edges.  Soon  saw 
another  on  bank  among  pickerel  weed.  Crawled  upon  it  on  my  knees  and 
caught  it;  it  squealed  as  some  frogs  will  do  when  caught.  The  children  went 
along  and  scared  another,  which  gave  a  startled  note  of  the  R.  catesbeiana 
order.  These  two  arc  the  largest  we  have  caught.  They  are  bigger  than  any 
green  frog.  The  transformed  ones  arc  somewhat  like  green  frogs  but  with 
more  intensified  spotting  on  belly.  These  as  transformed  and  adults  have  no 
costal  fold. 

June  9,  Callahan.  Went  out  last  night  at  bridge  which  is  type  locality  of 
JR.  hecl{scheri.  Saw  a  female  near  edge  of  Pontedena  bank.  Later  heard  a  big 
frog  jump  in.  When  I  approached  that  area,  found  a  male,  a  large  one,  on 
bank  and  among  Pontedena,  not  at  edge.  Saw  another  female.  Caught  her. 
Later  went  over  to  Mr.  Davis'  brick-yard  pools.  Many  transforming  frogs 
here  and  countless  tadpoles.  Sometimes  these  tadpoles  were  so  thick,  one  could 
reach  in  with  hand  and  catch  them.  On  the  east  side  saw  immense  school  of 
little  black  ones  with  the  yellowish  croisbands.  Amongst  them  are  some  bigger 
ones  about  %  inch  in  body.  This  means  at  least  two  lots  have  already  bred. 
This  is  the  first  lot  of  young  seen.  These  pools  have  water  lilies,  Sagittana, 
water  hyacinth,  Hydrocotyle,  and  many  other  plants. 

Authorities9  corner: 
M.  J.  Allen,  Miss.,  1932,  pp.  10-11. 
C.  S.  Brimley,  N.C.,  1941,  p.  3. 


FROGS:  RANIDAE  473 

Utah  Frog 

Rana  onca  Cope.  Plate  XCIX;  Map  34. 

Are  Rana  onca  and  Rana  fisheri  one?  Is  Rana  onca,  Rana  pipiens?  Many 
collections  in  Utah,  Nevada,  and  Arizona  are  needed  to  solve  this  possible 
R.  ptptens  derivative  if  it  be  such. 

See  L.  Stejneger,  Calif.,  1893,  p.  227,  under  R.  fishen. 

In  1940  J.  M.  Linsdale  re-established  Rana  fishen  Stejneger  and  retained 
R.  onca  Cope  as  distinct  from  R.  fishen.  Excerpts  from  this  author  follow: 

Rana  onca  "inhabits  streams  and  springs  in  southeastern  part  of  the  state; 
localities  are  along  the  Virgin  and  Lower  Muddy  rivers.  They  range  in  alti- 
tude from  about  1,200  feet  up  to  around  2,000  feet. 

"Recently  collected  series  of  specimens  from  several  localities  near  the 
Virgin  River  indicate  that  this  form  is  distinct  from  both  Rana  pipiens  and 
Rana  fishen.  From  the  former  it  differs  by  having  smaller  and  fewer  spots  and 
these  on  a  paler  background  and  with  less  distinct  gray  borders,  larger  tympa- 
num, and  shorter  legs.  It  is  nearer  to  fishen  which  reaches  the  extreme  de- 
velopment in  these  characters,  having  still  fewer  dorsal  spots  and  much  shorter 
hind  legs.  The  characters  of  smaller  si/e  and  absence  of  longitudinal  folds 
between  dorso-lateral  ridges  seem  not  to  be  diagnostic  in  the  Nevada  material" 
(J.  M.  Linsdale,  Nev.,  1940,  pp.  210-211;  see  also  his  comments  about 
R.  fishen}. 

"In  1928,  I  had  an  opportunity  of  examining  the  type  specimen  of  this 
species  described  by  Professor  Cope  in  1875.  For  a  long  time  this  species  was 
questionable  and  the  type  was  unknown;  as  a  result  Dr.  Stejneger  names  the 
specimens  taken  in  the  Vegas  Valley  by  members  of  the  Death  Valley  Ex- 
pedition Rana  fisheri.  The  rediscovery  of  the  type  specimen  which  was  so 
faithfully  reproduced  in  1875  places  R.  fishen  as  a  synonym"  (V.  M.  Tanner, 
Utah,  1931,  pp.  189-190). 

Range:  Along  the  Virgin  River,  southeastern  Nevada,  northwestern  Ari- 
zona, and  southwestern  Utah. 

Size:  Linsdale  collected  78  specimens.  These  measure  as  follows:  adults 
44-84  mm.,  males  44-68;  females  51-84  mm.  Rodgers  took  males  of  44.0-55.5 
mm.  and  females  of  48.5-82.0  mm. 

General  appearance:  Notes  on  Linsdale's,  Rodger's,  Smiley 's,  and  Hall's 
specimens  follow : 

1.  Light  line  from  below  eye  and  under  tympana  to  behind  angle  of  mouth 
is  prominent,  this  being  absent  in  adults  of  R.  fishen.  What  is  it  in  the  leg 
that  makes  it  possible  for  heel  of  R.  onca  to  go  between  eye  and  snout?  Is  it 
the  femur?  Dark  spot  from  eye  to  nostril  common  in  R.  onca. 

2.  MVZ  no.  19608,  47.5  mm.,  male,  April  25,  1936.  Seems  to  have  tail  apron 
long  after  transformation. 

3.  No.  20433,  47.5  mm.,  male,  May  3,  1956.  Big  thumb. 


474  HANDBOOK  OF  FROGS  AND  TOADS 

4.  No.  20662,  47  mm.,  male,  June  10,  1936.  Thumb  not  so  well  developed. 
Is  it  of  breeding  size? 

5.  No.  20660,  47.5  mm.,  male,  June  10, 1936.  Thumb  not  so  well  developed. 
Is  it  past  breeding? 

6.  No.  19582, 48.5  mm.,  female,  April  25, 1936.  Is  it  breeding? 

7.  No.  13888, 52  mm.,  male,  Sept.  30, 1931.  Thumb  pads  not  well  developed. 

8.  No.  16571,  60  mm.,  female,  April  25,  1936.  Ripe. 

Color:  Spring  2  miles  southeast  of  Overton,  Nev.,  May  18,  1942.  The  gen- 
eral impression  is  brown  in  the  forward  dorsum  and  gray  in  the  rear  and  upper 
forearms.  The  dorsal  background  color  of  top  of  head  from  front  of  snout, 
between  nostrils,  m  front  of  eye,  and  upper  eyelid  is  clear  sudan  brown  or 
buffy  brown.  The  tympanum  is  the  same  with  a  spot  of  cartridge  buff  or 
ivory  yellow  in  the  middle.  The  background  color  rearward  becomes  drab; 
rear  of  legs  is  smoky  gray,  forelegs  are  pale  ecru-drab  or  ecru-drab  on 
brachium.  The  sides  are  intermediate  between  color  of  hind  legs  and  mid- 
dorsum,  with  two  lateral  spots  touching  the  dorsolateral  folds  and  three  more 
lower  on  the  side.  The  costal  folds  are  low,  are  tilleul  buff  or  pale  pinkish 
cinnamon.  Between  the  folds  are  two  irregular  rows  of  dark  spots  with  one 
spot  between  in  middorsum  and  one  extra  in  pelvic  region.  There  is  a  touch 
of  peacock  green  in  middorsum  and  on  outer  edges  of  spots.  The  spots  are 
cinnamon-brown  or  light  brownish  olive  with  tendency  toward  smoky  gray 
margins.  The  bars  on  hind  legs  are  light  grayish  olive  in  the  center  and  grayish 
olive  on  margins.  There  is  a  little  spot  on  heel,  three  bars  on  tibia,  a  prominent 
spot  near  femur-tibia  articulation,  and  two  to  three  bars  on  the  femur.  On 
fore  limb  there  is  an  oblique  band  on  antebrachmm,  one  on  elbow,  and  a 
prominent  oblique  brown  bar  from  insertion  halfway  down  on  front  of 
brachium.  From  front  of  nostril  to  eye  is  a  band  of  light  brownish  olive  or 
cinnamon-brown.  The  margins  of  upper  and  lower  jaws  are  smoky  gray. 
There  is  a  thin  line  of  peacock  green  under  the  eye,  extending  forward  be- 
tween the  light  brownish  olive  band  above  and  the  smoky  gray  upper  labial 
region.  From  under  the  eye  extending  backward  under  the  tympanum  and 
downward  on  shoulder  to  arm  insertion  is  a  prominent  light  stripe  like  the 
dorsolateral  folds.  The  under  surfaces  are  white  with  the  groin  somewhat 
spotted  and  touched  with  yellow,  which  color  is  also  toward  the  front.  The 
iris  is  black  with  flecks  and  bands  of  orange-cinnamon. 

Structure:  "Rana  onca  Cope  sp.  nov.  .  .  .  Head  oval,  muzzle  sloping  to 
the  lip.  Diameter  of  tympanic  membrane  equal  distance  between  nares  and 
between  nostril  and  orbit,  and  three-fourths  the  diameter  of  the  orbit  or  the 
distance  from  nares  to  margin  of  lip  in  front.  Vomerine  teeth  in  vascicuh  be- 
hind the  line  connecting  the  posterior  borders  of  the  choanae.  A  dermal  fold 
on  each  side  of  the  back,  and  a  short  one  behind  the  angle  of  the  mouth,  with 
some  scattered  warts  on  the  sides;  skin  otherwise  entirely  smooth.  Toes  ob- 
tuse, with  wide  webs  reaching  to  the  base  of  the  penultimate  phalange.  One 


FROGS:  RANIDAE  475 

long  metatarsal  tubercle;  no  fold  on  the  tarsus;  a  dermal  border  on  outer  toe. 
The  heel  extends  beyond  the  end  of  the  muzzle. 

"Light  brown  above;  below  yellow.  Three  rows  of  rather  distant  solid 
black  spots  between  the  dorsal  folds;  two  or  three  rows  on  each  side;  none 
of  the  spots  yellow-bordered.  Head  unspotted;  no  band  on  the  lip.  A  brown 
vertical  band  on  the  front  of  the  humerus.  Scattered  spots  on  tibia  and  femur; 
clouded  spots  on  the  posterior  face  of  the  femur.  Size  of  Rana  clamata. 

"This  frog  of  which  a  female  specimen  was  obtained,  combines  characteris- 
tics of  different  groups;  its  coloration  resembles  somewhat  that  of  the  eastern 
or  typical  form  of  Rana  halecina,  but  the  full  palmation  of  the  hind  foot  is 
that  of  R.  montezumae  and  R.  catesbeiana.  It  is  also  quite  similar  to  the 
variety  of  R.  temporana  from  California,  called  R.  dray  torn  by  Baird  and 
Girard  and  R.  longipes  by  Hallowell.  The  feet  are  shorter,  the  hind  feet  be- 
ing twice  as  long  as  the  head  to  the  posterior  border  of  the  tympanum,  while 
in  R.  t.  draytom  it  is  2.5  times  as  long.  The  R.  onca  lacks  the  black  cheek-patch 
of  the  jR.  temporana.  Specimen  secured  in  1872  in  Utah  by  Dr.  H.  C.  Yarrow. 
The  plate  affords  view  ot  the  dorsum  and  mouth  of  this  species"  (H.  C. 
Yarrow,  Ariz.,  1875,  pp.  528-529). 

This  frog  in  snout  and  general  appearance  looks  more  like  R.  sphenocephala 
than  R.  pipiens.  It  is  of  R.  pipiens  class  but  smaller. 

"This  rare  Utah  species  may  be  readily  separated  from  all  other  native 
Batrachians  except  R.  ptptens  and  from  this  species  it  differs  in  having  no 
definite  narrow  longitudinal  folds  between  the  dorsolateral  ridges,  its  smaller 
size,  and  habitat  in  the  extreme  southwestern  corner  of  Utah  along  the  Virgin 
River.  The  head  is  as  broad  as  long  and  depressed;  the  nostrils  are  nearer  the 
eyes  than  the  end  of  the  snout;  canthus  rostralis  almost  absent  in  the  specimens 
before  me.  Tympanic  membrane  large  and  smooth.  The  limbs  are  well  de- 
veloped, digits  long  and  webs  well  formed.  Skin  above  smooth  with  a  few 
warts  and  tubercles  about  the  head  and  neck.  The  general  color  above  is  brown 
to  olive  green  with  spots  that  arc  bordered  with  grey.  The  noticeable  dorso- 
lateral fold  is  brown  in  the  specimens  examined.  The  undersurface  is  white 
over  the  belly  and  yellowish  on  the  legs"  (V.  M.  Tanner,  Utah,  1951,  p.  189). 

Voice:  Its  call  is  not  so  long  or  loud  as  the  call  of  the  bigger  R.  pipiens. 

Breeding:  Most  of  the  captures  were  late  April.  This  is  doubtless  the  height 
of  its  breeding  season.  It  may  breed  March  to  May.  Kggs  and  tadpoles  are 
not  of  record.  Transformation  probably  is  from  25  to  30  mm.  or  more  in  June. 
Were  the  specimens  secured  April  25,  19^6,  at  Overtoil,  sizes  36  to  42  mm., 
from  the  year  before ? 

Journal  notes:  May  18,  1942,  Glcndalc.  Went  to  the  ranger  of  Boulder  Dam 
Museum.  He  said  that  "2  miles  S.  E.  of  Overton,"  where  Lmsdale  and  Rodgcrs 
caught  the  frog,  must  be  a  spring  seep  ]*>  mile  beyond  where  the  CCC  boys 
were  making  a  diversion  dam  on  Muddy  River.  The  museum  is  %  niile  from 
Overton. 


476  HANDBOOK  OF  FROGS  AND  TOADS 

Went  to  seepage  area,  quite  dry.  Saw  a  frog  leap  into  a  clump  of  grass, 
missed  it.  My  impression  was  gray.  Went  two  steps  farther  and  another  (prob- 
ably the  same)  leaped  into  brush.  I  whipped  him  out  and  he  leaped  into  a  pool. 
Got  it.  Later,  lost  another  which  leaped  into  rushes. 

Beaver  Dam  Lodge,  Ariz.  We  heard  a  low  short  croak — the  frog  we  wanted. 
After  a  long  effort  I  located  it  in  the  tules,  but  with  one  flashlight  on  right 
wrist  and  one  in  my  hand  and  with  a  net  and  a  bag  at  my  waist,  I  missed  in 
my  grab  at  him.  He  looked  like  the  one  we  captured  at  Overton. 

Authorities9  corner:  "A  search  of  several  years,  for  this  species,  finally  ended 
successfully  when  two  specimens  were  taken  in  June,  1928,  in  a  small  stream 
south  of  St.  George.  While  Dr.  Yarrow  does  not  give  the  exact  locality  for 
this  species  yet  from  his  itinerary  and  subsequent  collections  we  are  reasonably 
sure  in  concluding  that  he  captured  his  specimens  somewhere  along  the  Virgin 
River  in  Washington  County  [Utah|"  (V.  M.  Tanner,  Utah,  1929,  p.  49). 

"This  species  was  first  observed  at  the  mouth  of  Boulder  Wash,  Colorado 
River,  where  the  rising  waters  of  the  reservoir  had  submerged  a  small  cat-tail 
swamp.  Two  specimens  were  seen  clambering  about  on  small  bits  of  drift- 
wood a  few  feet  from  the  bank.  One  specimen  was  collected  at  Littlefield, 
Arizona,  and  five  specimens  were  taken  on  the  Muddy  River  at  Glendale, 
Nevada.  Although  Woodbury  (1930)  has  recorded  R.  pipiens  from  Zion  Can- 
yon on  the  Mukunteweap  River,  a  tributary  of  the  Virgin,  it  would  seem 
improbable  that  two  such  closely  related  species  as  onca  and  pipiens  would  oc- 
cur in  contiguous  regions  of  the  same  drainage  system.  More  recently,  Eaton 
has  stated  that  specimens  collected  further  eastward  in  Utah  'are  intermediate' 
and  he  doubts  the  validity  of  onca.  It  is  more  probable  that,  if  not  valid  species, 
the  two  forms  may  be  subspecific"  (R.  B.  Cowles  and  C.  M.  Bogert,  Nev., 
1936,  p.  36). 

Pickerel  Frog,  Marsh  Frog,  Tiger  Frog  (DeKay),  Leopard  Frog  (DeKay), 
Cold  Swamp  Frog,  Swamp  Frog,  Spring  Frog,  Le  Conte's  Leopard  Frog 

Rana  palustris  Le  Conte.  Plate  C;  Map  32. 

Range:  From  central  plains  to  Atlantic  seaboard;  from  the  Gulf  States  to 
Hudson  Bay. 

"Rana  palustris  was  not  found  (Northern  Pontiac  Co.,  Quebec)  although  it 
has  been  taken  in  the  Laurentian  Mountains  (Lat.  46°N.).  Cope  (1889)  as- 
signed three  specimens  collected  by  Drexler  from  'James  Bay,  N.B.'  to 
palustris.  The  identification  was  confirmed  by  Preble  (1902).  The  locality 
designated  has  been  assumed  by  later  authors  to  be  James  Bay  proper,  and 
since  there  is  no  James  Bay  in  New  Brunswick  this  is  probably  the  correct 
interpretation.  Since  there  are  no  other  records  of  this  species  from  high  lati- 
tudes, Cope's  record  should  not  be  accepted  without  confirmation"  (R.  Grant, 
Que.,  1941,  p.  153). 


FROGS:  RANTt>AE 


477 


^ 


Plate  XCIX.  Rana  onca.  (X%)-  From 
2  miles  southeast  of  Overton,  Nev. 


Plate  C.  Rana  palustris.  1,3.  Males  (X%) 
2.  Male  croaking  (X%)«  4-  Female  (X%) 


478  HANDBOOK  OF  FROGS  AND  TOADS 

Habitat:  Sphagnum  bogs,  marl  ponds,  cold  streams,  shallows  of  mill  ponds, 
quiet  water  of  bayous,  rocky  ravines.  It  is  the  most  abundant  frog  in  the  rock- 
bottomed  and  rock-walled  ravines  of  central  New  York,  where  it  quite  fre- 
quently breeds  in  the  deep  holes  at  the  bases  of  waterfalls.  It  is  also  found  in 
the  sources  of  clear  trout  streams.  After  breeding  time,  the  frog  is  found  in 
meadowlands.  Some  individuals  remain  in  ravines  and  similar  places.  They 
hibernate  in  water. 

"The  pickerel  frog  is  usually  restricted  to  flowing  streams  either  cool  or 
warm.  Specimens  have  been  taken  in  trout  brooks,  at  55°F.  and  in  drainage 
creeks,  at  8o°F.  In  Glengarry  County  in  late  September  we  found  specimens 
a  considerable  distance  from  any  water"  (G.  C.  Toner  and  N.  dc  St.  Remy, 
Ont.,  1941,  pp.  13-14). 

Size:  Adults,  1^5-5%  inches.  Males,  46-64  mm.  Females,  49-79  mm. 

General  appearance:  The  pickerel  frog  at  first  glance  looks  much  like  the 
meadow  frog.  It  is  slender,  smooth-skinned,  medium  in  size,  usually  brown 
in  color,  with  light  raised  ndges  extending  backward  from  the  eyes.  The  dark 
spots  arc,  however,  square  in  shape  and  dark-bordered,  regularly  arranged  in 
two  rows  with  an  occasional  third  spot  between  the  dorsolateral  folds,  and 
covering  more  than  one-half  of  that  area.  The  spots  on  the  sides  also  are  square 
and  arranged  regularly,  the  upper  row  close  to  the  dorsolateral  fold.  There 
is  a  dark  streak  from  the  eye  to  the  nostril.  Underneath  it  is  glistening  white 
on  the  forward  part  and  yellow  or  orange  on  the  underside  of  the  legs,  in  the 
groin,  forward  along  the  sides  of  the  belly,  and  on  the  underside  of  forearms. 
The  legs  arc  barred  with  dark  bands. 

Color:  Male.  McLean,  N.Y.,  June  2, 1929.  Ground  color  from  back  of  snout 
to  vent  and  between  dorsolateral  folds,  ecru-olive,  also  same  on  interspaces  of 
tibia  and  between  crossbars  of  femur.  Interspaces  of  forelegs  and  sides  below 
dorsolateral  folds  avcllaneous  or  vmaceous-buff.  Snout  and  line  over  eye  same 
or  tawny  olive.  Dorsolateral  fold  citron  yellow  or  ochiaccous-huff,  at  times 
capucinc  yellow.  Median  spot  ahead  of  eye,  eyelid  spot,  stripe  eye  to  snout, 
two  rows  down  back,  regular  and  smaller  lower  side  spots,  crossbars  on  legs, 
etc.,  brownish  olive  or  black  with  lighter  dots  of  light  yellowish  olive  which 
give  tibial  crossbars  and  some  others  a  rather  "greenish"  cast.  Under  parts 
of  hind  legs  and  forelegs,  groin,  and  lower  belly  orange  or  cadmium  yellow, 
becoming  a  narrow  light  orange-yellow  band  along  side  to  axilla.  Rear  of 
femur  raw  sienna  or  aniline  yellow.  Rest  of  under  parts  cream  color.  Tympa- 
num wood  brown.  Ins  black  or  clove  brown  dotted  below  with  light  russet- 
vinaceous  flecks.  Above  pupil  is  a  line  of  light  green-yellow  or  empire  yellow. 
Edge  of  eyelid  with  a  little  of  neva  green.  Thumb  pad  blackish  brown  (i). 
Stripe  on  upper  jaw  extending  almost  to  arm  insertion  is  honey  yellow  at  front, 
becoming  cartridge  buff  at  the  rear,  edged  below  by  brownish  olive.  Promi- 
nent blackish  brown  (i)  longitudinal  spot  at  base  of  arm  and  in  front. 


FROGS:  RANIDAE 


479 


Female.  McLean,  July  21,  1929.  Ground  color  of  back  from  snout  to  vent 
and  the  interspaces  on  tore  and  hind  legs  deep  olive-huff  or  grayish  olive. 
Below  the  dorsolateral  folds  and  on  the  interspaces  of  forearms  and  hind  legs 
the  frogs  have  a  tendency  to  become  light  drab  or  cinnamon-drab.  The  upper 
eyelid  is  vinaceous-fawn.  The  dorsolateral  fold  is  light  ochraceous-salmon  or 
cinnamon-buff.  One  median  spot  ahead  of  the  eye,  eyelid  spot,  stripe  from  eye 
to  snout,  the  two  rows  down  the  back,  and  a  smaller  row  below  the  dorso- 
lateral fold  are  buffy  olive.  So  also  the  bars  on  the  legs;  the  bars  on  the  hind 
legs  and  the  spots  below  the  dorsolateral  fold  may  be  light  brownish  olive. 
The  vermiculations  on  the  rear  of  thighs  and  the  margins  to  the  spots,  the 
oblique  bar  on  the  arm  insertion,  and  the  oblique  stripe  over  tympanum  from 
eye  to  arm  may  be  black,  olive,  or  brownish  olive.  As  in  the  males,  all  of  the 
dorsal  spots  and  leg  bars  may  have  a  greenish  cast  because  of  yellowish  olive 
or  light  yellowish  olive  dots  in  their  centers.  In  the  groin  and  axilla  is  cadmium 
yellow,  but  the  rear  half  of  underside  of  femur,  the  whole  underside  of  tibia, 
and  the  inner  side  of  leg  are  raw  sienna  or  mars  yellow. 

This  is  also  true  of  vermiculations  on  the  rear  of  femur.  The  rest  of  under 
parts,  including  some  of  the  middle  of  the  lower  belly  and  some  of  basal  half 
of  femur,  are  cream-colored.  The  tympanum  is  wood  brown.  Stripe  on  jaw 
is  cartridge  buff  with  brownish  olive  edge  below  it.  Eye  as  in  the  male. 

Structure:  Thumb  much  enlarged  in  breeding  males;  vocal  sacs  of  males 
small  lateral  swellings  between  the  tympanum  and  the  arm;  the  skin  has 
strong  acrid  secretion,  irritating  to  the  mouth  of  the  dog  who  tries  to  eat  one, 
a  secretion  which  will  kill  other  frogs  put  in  the  same  jar  of  water. 

"Rana  pahtstrts,  or  marsh  frog;  colour  above  pale  brown,  with  two  longi- 
tudinal rows  of  dark  brown  spots  on  the  back,  and  the  same  number  on  the 
sides,  hind  part  of  the  thighs  yellow,  spotted  with  black"  (J.  Le  Conte,  Gen., 
1825,  p.  282). 

Voice:  It  has  a  low-pitched  grating  cioak  with  little  carrying  power,  shorter 
and  higher  than  Rana  ptptens,  and  more  prolonged  and  lower  than  Rana 
sylvattca.  These  males  frequently  croak  beneath  the  surface  of  the  water,  while 
in  the  embrace.  Of  our  local  frogs,  this  form  and  the  rather  silent  wood  frog 
make  the  least  audible  disturbance  at  the  breeding  season.  The  call  is  so  char- 
acterless that  the  tendency  is  to  link  it  with  a  subdued  Rana  pipicns,  but  the 
latter  has  not  so  short  a  note.  The  males  occasionally  answer  each  other  with 
a  croak  totally  different  from  the  normal  one.  E.  D.  Cope  (Gen.,  1889,  p.  408) 
says,  "Its  note  is  a  low  prolonged  croak,  somewhat  resembling  the  sound  pro- 
duced by  tearing  some  coarse  material." 

Breeding:  These  frogs  breed  from  April  23  to  May  15.  They  gather  in  large 
numbers,  and  often  in  a  small  area  6  feet  square  or  less  one  finds  12  to  15 
pickerel  frogs  mating  or  pairs  laying.  The  brown  and  bright  yellow  eggs  are 
submerged  attached  to  twigs  or  grass  stems,  form  a  firm  globular  mass  3/^-4 


480  HANDBOOK  OF  FROGS  AND  TOADS 

inches  (87-100  mm.)  in  diameter,  and  number  2000-3000.  The  egg  is  Mu 
inch  (1.6  mm.)  in  diameter,  envelopes  l/g  inch  (2.8  mm.)  and  %  inch  (4  mm.), 
a  little  smaller  than  those  of  Rana  pipiens.  The  tadpole  is  large,  3  inches 
(75.8  mm.),  greenish  in  color,  the  body  and  tail  covered  with  fine  black  dots, 
the  tail  crests  black  or  very  clouded,  and  the  belly  cream.  The  tooth  ridges 
are  %,  %.  After  70-80  days,  the  tadpoles  transform  in  August  at  %-iVia 
inches  (19-27  mm.). 

The  species  usually  seeks  the  shallows  for  egg  laying,  though  almost  every 
year  we  find  some  egg  masses  in  the  middle  of  ponds  where  the  water  is  1-3 
feet  deep.  Like  Rana  sylvatica  and  Rana  pipiens,  it  frequently  lays  m  special 
areas.  In  1907  in  an  area  3  by  3  feet,  18  bunches  were  deposited;  in  another 
12  bunches  were  recorded.  In  1911,  in  another  spot  3  by  3  feet,  31  bunches  were 
found.  All  these  areas  gave  excellent  illustration  of  the  building  of  bunches, 
one  upon  another.  In  one  case  four  successive  bunches  upon  one  twig  were  re- 
corded. The  greatest  number  on  one  support  was  secured  May  5,  1909,  when 
seven  were  noted.  Quite  frequently,  on  tufts  of  grass  or  sticks  one  finds  a 
bunch  or  more  of  spotted  salamander  eggs  or  meadow-frog  eggs  with  one  of 
two  pickerel  frog  masses  immediately  above  them,  seldom  below,  because 
laid  later. 

Journal  notes:  April  28,  1911,  Ithaca,  N.Y.  When  I  approached  the  Rana 
palustris  egg  area  (just  discovered  today)  there  were  within  an  area  of  3  feet 
square  5  mated  pairs  and  21  unmated  males.  Of  the  mated  pairs,  the  males 
were  much  lighter  than  the  females.  A  female  will  back  up  to  a  stem  and 
clasp  it  with  hind  legs,  then  change  position  or  back  to  another  stem.  Some- 
times this  will  be  a  stem  with  one  to  three  or  four  egg  masses  on  it  already. 
Her  nervous  leg  movements  and  many  changes  often  loosen  or  fray  the  former 
egg  masses.  In  the  area  mentioned  above  there  were  already  eight  egg  masses. 
Often  when  a  pair  took  a  position  other  males  interfered.  In  such  cases  the 
female  helped  her  consort  to  kick  them  away.  While  in  the  embrace,  the  male 
frequently  gave  his  low  grating  note.  Occasionally  a  pair  put  up  for  air. 
Finally  one  pair  began  to  lay.  She  was  perfectly  horizontal  and  flattened,  her 
hind  legs  drawn  up,  the  heels  together,  her  front  feet  together;  with  his  hind 
legs  slightly  within  hers,  he  seemed  to  help  push  the  eggs  back.  This  egg 
laying  occupied  about  3  minutes  in  all,  with  about  10  to  12  fertilizations.  I 
left  the  area  after  10:50,  returning  at  11:00,  and  found  another  mass  had^been 
laid. 

Authorities9  corner: 

D.  H.  Storer,  Mass.,  1839,  p.  238.  J.  A.  Weber,  N.Y.,  1928,  p.  109. 

H.  C.  Bumpus,  R.I.,  1886,  pp.  7-8.  H.  M.  Smith  and  B.  C.  Brown,  Tex., 

G.  M.  Allen,  N.H.,  1899,  p.  69.  1947,  pp.  47-50. 

Anita  Daugherty's  unpublished  thesis  (University  of  Rochester)  may  have 
pertinence  in  the  understanding  of  the  Rana  pipiens  complex, 


FROGS:  RANIDAE  481 

Meadow  Frog,  Leopard  Frog,  Shad  Frog,  Herring  Hoppers,  Common  Frog, 
Berlandier's  Frog,  Spotted  Frog,  Grass  Frog,  Water  Frog,  Peeping  Frog, 

Olive-colored  Frog 

Rana  pipiens  pipiens  Schreber.  Plate  CI;  Maps  33-34. 

Range:  Widespread  and  common  over  North  America,  east  of  the  eastern 
edge  of  the  Pacific  Coast  states,  from  the  extreme  north  into  Mexico. 

Specifically  on  the  west  it  extends  from  Mexico  to  the  southeastern  corner 
of  California,  up  the  Colorado  River  to  Nevada,  then  to  northeastern  Cali- 
fornia, eastern  edges  of  Oregon,  Washington,  and  British. Columbia  to  north- 
ern Alberta  or  even  to  Mackenzie,  thence  southeastward  to  Laurent ian  Moun- 
tains and  Gaspe  Peninsula,  Labrador,  and  Newfoundland;  thence  along  coast 
to  northern  New  Jersey  and  along  southeastern  states  in  the  Piedmont  and 
the  mountains  mainly  above  the  Fall  line,  i.e.,  western  two-thirds  of  Virginia, 
western  half  of  North  Carolina,  western  South  Carolina,  northern  Georgia 
and  Alabama,  eastern  Tennessee  and  Kentucky;  thence  westward  through 
northern  Indiana,  Illinois,  and  Missouri  to  Kansas  and  south  through  the 
Panhandle  of  Texas  to  extreme  southern  Texas  and  Mexico. 

Habitat:  In  spring  in  swampy  marsh  lands,  upland  backwaters,  overflows, 
and  ponds.  In  summer  in  swamp  lands,  grassy  woodland,  or  hay  or  grain 
fields  in  cultivated  districts.  They  spend  the  winter  in  pools  or  marshes. 

Size:  Adults,  21/ly-41/i^  inches.  Males,  52-80  mm.  Females,  52-102  mm. 

General  appearance:  These  frogs  are  slender  in  form,  smooth-skinned,  me- 
dium in  size,  brown  or  green  in  general  color,  with  two  light  raised  stripes 
extending  backward  from  the  eye.  Between  these  dorsolateral  folds  are  two 
or  three  rows  of  irregularly  arranged,  rounded  dark  spots  with  light  borders. 
On  the  sides  are  more  rounded  dark  spots  with  light  borders,  irregularly  ar- 
ranged in  three  or  four  rows.  There  is  usually  a  dark  spot  on  the  top  of  either 
eyelid.  There  is  a  light  line  along  the  jaw,  below  the  ear,  and  over  the  arm, 
bordered  below  with  a  dark  stripe.  The  legs  are  barred  with  light-bordered 
dark  bands.  Beneath,  it  is  a  glistening  white  to  yellow  or  orange  under  legs 
m  Arizona.  (See  R.  pipiens  complex.) 

Color:  Male.  Ithaca,  N.Y.,  from  Harry  Leighton,  July  29,  1929.  The  back 
is  Saccardo's  olive  to  buffy  brown  with  spots  on  back  and  sides  of  chestnut 
brown  or  olive-brown  with  black  rims.  The  spots  are  surrounded  by  rings  of 
apple  green  to  yellow-green,  which  is  the  color  extending  from  nostril  to  eye, 
over  tympanum,  and  along  the  side  just  below  the  dorsolateral  fold.  There 
is  a  pale  vinaceous-fawn  to  cartridge  buff  line  along  upper  jaw  to  above  arm 
insertion,  bordered  below  by  olive-brown,  which  in  turn  is  margined  on  lower 
edge  by  cartridge  buff  or  white.  The  dorsolateral  folds  are  pale  vinaceous- 
fawn  to  pale  cinnamon-pink.  The  interspaces  on  forearms  and  lower  sides 
are  light  vinaceous-fawn  to  ecru-drab.  There  is  an  olive-brown  dash  on  front 
of  brachial  insertion.  Upper  parts  of  hind  limbs  are  like  the  back,  with  inter- 


482  HANDBOOK  OF  FROGS  AND  TOADS 

spaces  on  rear  of  femur  viridme  yellow  or  neuvider  green  spotted  olive-brown 
to  black.  In  the  groin  is  a  little  bright  green-yellow.  The  tympanum  is 
vmaceous-buff  or  cinnamon  with  spotting  of  olive-brown.  Under  parts  are 
white  or  cartridge  buff.  The  iris  is  black  or  olive-brown  with  an  outside  half- 
moon  back  and  front  of  neuvider  green;  the  lower  part  of  ins  and  pupil  rim 
are  spotted  with  vinaceous  buff  and  with  a  longitudinal  band  over  the  pupil 
and  pupil  rim  of  light  greenish  yellow.  The  thumb  has  basal  pad  of  hair  brown 
or  dark  grayish  olive. 

Female.  Ithaca,  N.Y.,  July  23,  1950.  The  dorsolateral  folds,  canthus  rostrahs, 
rear  of  upper  eyelid,  and  supplementary  folds  between  the  dorsal  folds  are 
mikado  brown  to  wood  brown  or  avellaneous.  The  latter  colors  are  the  back- 
ground of  back,  sides,  and  forelegs.  On  rear  legs  there  is  a  little  of  this  color 
on  the  distal  cephalic  portion  of  femur,  on  dorsal  tibia,  and  on  foot.  The  dorsal 
spots  are  Prout's  brown  or  possibly  bister.  The  borders  of  spots  are  neva  green 
or  calliste  green.  The  lower  lateral  spots  and  spots  of  rear  of  femur  are  bone 
brown  without  light  borders.  The  interstices  of  the  groin  and  rear  of  femur 
are  neuvider  green.  Some  of  the  lower  lateral  spots  and  the  spots  on  the 
arm  and  sometimes  on  foot  are  tillcul  buff.  The  light  stripe  cm  upper  jaw  to 
arm  insertion  is  tilleul  buff  becoming  olive-buff  or  pale  olive-buff  over  arm 
insertion.  The  tympanum  is  same  color  as  dorsal  folds.  The  iris  is  like  the 
male's.  Under  parts  are  white. 

Structure:  Lateral  folds  prominent;  head  shorter  than  R.  p.  sphenocephala; 
snout  medium;  vocal  sacs  of  males,  lateral,  between  tympanum  and  arm, 
swelling  out  as  round  balls  over  the  arms,  with  extensions  down  the  sides; 
more  spots  below  dorsolateral  fold  than  in  Rana  p.  sphenocephala. 

Voice:  The  croak  is  a  long,  low  guttural  note,  3  or  more  seconds  long, 
followed  by  three  to  six  short  notes  each  a  second  or  less  in  length,  or  the 
short  notes  may  precede  or  be  interspersed.  In  early  spring  the  leopard  frogs 
form  the  second  swamp  chorus  to  arrest  attention.  Though  not  possessing 
loud  voices  individually,  their  concourses  in  our  swamps,  when  most  vocif- 
erous, cannot  pass  unnoticed.  Usually  the  call  is  heard  from  males  at  the 
surface  of  the  water,  and  before  they  are  mated.  Occasionally,  when  wading 
through  an  egg  area,  one  hears  croaks  which  at  first  puzzle;  they  come  from 
the  mated  and  mating  frogs  beneath  the  water  and  often  reveal  the  game  on 
the  bottom. 

Breeding:  This  species  breeds  from  April  i  to  May  15.  The  egg  mass  is  a 
flattened  sphere,  3-6  inches  (75-150  mm.)  by  2-3  inches  (5-75  mm.).  The 
eggs  are  black  and  white,  Vi «•>  inch  (1.6  mm.)  in  diameter,  the  envelopes  %  inch 
(3.4  mm.)  and  H  inch  (5  mm.).  The  tadpole  is  large,  3%  inches  (84  mm.), 
the  tail  lighter  than  the  body,  its  crests  translucent  marked  with  fine  spots 
and  pencilings.  The  tooth  ridges  are  %,  %.  After  60  to  80  days,  the  tadpoles 
transform  in  July  at  %-\V±  inches  (18-31  mm.). 

These  frogs  have  a  tendency  to  congregate  in  large  numbers  at  the  breeding 


FROGS:  RANIDAE  483 

time  and  often  40  or  more  bunches  of  eggs  are  recorded  within  a  small  area. 
The  egg  masses  are  plinthlike  and  attached  to  submerged  cattails,  twigs, 
sticks,  or  grass,  or  even  rest  on  the  bottom  unattached.  They  occur  in  the  open, 
unprotected,  marshy  expanses,  or  in  overflows  where  edges  and  bottoms  have 
plenty  of  grass.  Many  of  the  egg  masses  are  laid  in  very  shallow  water,  and 
it  does  not  require  very  much  evaporation  to  leave  them  high  and  dry. 

Journal  notes:  April  n,  1907,  Iihaca,  N.Y.  In  Bool's  backwater,  I  found  one 
bunch  of  Rana  pipiens  eggs.  Just  before  I  found  them  I  heard  a  croak  under 
my  feet.  There  were  two  frogs  trying  to  escape,  a  male  and  a  female.  .  .  .  The 
dead-end  stream  running  east  furnished  some  good  notes.  I  heard  several 
croaks  from  frogs  at  the  surface.  It  is  a  low  croak.  I  captured  a  male  Rana 
pipiens.  Around  were  eight  bunches  of  eggs.  Rana  pipiens  eggs  are  not  in  the 
spherical  masses  that  we  find  with  Rana  sylvatica*  They  are  more  flat;  the 
longest  diameter  may  be  5  or  6  inches,  but  seldom  is  the  other  diameter  more 
than  i%  or  2  inches.  Rana  pipiens  usually  seeks  places  where  the  edges  are 
grassy. 

April  25, 1911,  Dwyer's  Pond,  Ithaca,  N.Y.  Surface  temperature  56°,  bottom 
temperature  48°.  I  found  several  new  bunches  of  Rana  pipiens  eggs,  countless 
old  bunches,  immense  Rana  pipiens  areas  of  25-40  bunches,  and  several  Rana 
pipiens  croaking. 

Authorities'  corner: 

P.  Kalm,  Gen.,  1771,  II,  88-90.  B.  W.  Evermann  and  H.  W.  Clark, 

G.  H.  Loskiel,  Gen.,  1794,  p.  89.  N.Y.,  1920,  p.  638. 

W.  T.  Davis,  N.Y.,  1898,  p.  4. 

Plain  Meadow  Frog,  Burns'  Meadow  Frog,  Unspotted  Meadow  Frog 

[Mutant]  Rana  pipiens  burnsi  Weed.  Plate  CII;  Map  34. 

Range:  "The  species  seems  to  be  confined  to  northern  Iowa  and  southern 
Minnesota,  with  possibly  some  stragglers  in  western  Illinois  and  Wisconsin. 
.  .  .  This  specimen  (the  type)  and  twenty  paratypes,  were  received  from  New 
London,  Kandiyohi  County,  Minnesota.  There  are  about  eighty  other  speci- 
mens of  this  species  in  the  collections  of  Field  Museum,  from  Spicer,  Kandi- 
yohi County,  Minnesota;  Okabena,  Jackson  County,  Minnesota;  Rothsay, 
Wilkin  County,  Minnesota;  and  Astoria,  Deuel  County,  South  Dakota"  (A. 
C.  Weed,  Minn.,  1922,  pp.  108-109). 

Habitat:  Swampy  stretches  and  meadows.  (See  "Journal  notes"  for  this 
form  and  Rana  p.  fandiyohi.) 

Size:  "In  size  it  is  very  little  smaller  than  Rana  pipiens.  That  is,  the  very 
largest  individuals  of  the  latter  species  are  a  little  larger  than  the  very  largest 
ones  of  the  former"  (A.  C.  Weed,  Minn.,  1922,  p.  108). 

General  appearance:  These  frogs  are  built  like  meadow  frogs.  They  have 
the  long  snout  of  R.  p.  sphenocephala.  Their  general  color  and  first  appearance 


HANDBOOK  OF  FROGS  AND  TOADS 


Plate  C7.  Kana  pipiens  pipiens.  i.  Female 
(X1^).  2,5,  Males  (X%).  3*  Egg  masses  on 
the  bottom  of  a  pond  (X^)-  4-  ^gg  massi 
plinth  shaped 


Plate  CIL  Rana  pipiens  (burnsi)  (X%). 
1-6.  Males. 


FROGS:  RANIDAE  485 

is  like  a  bronzy  wood  frog  (R.  sylvatica)  without  the  black  mask.  The  costal 
folds  are  long,  prominently  raised,  and  frequently  a  light  buff  color.  Some 
frogs  are  apple  green,  some  wood  brown.  They  have  considerable  green  in 
axil  and  groin,  and  on  rear  of  femur.  Some  have  prominent  black  dash  on 
arm  insertion  and  black  spot  on  elbow,  and  prominent  black  band  on  canthus 
rostralis.  They  are  glistening  white  below.  They  are  beautiful  frogs  and  re- 
mind one  of  wood  frogs. 

Aug.  24,  1930,  New  London,  Minn.  We  took  two  large  males  on  Lower 
Florida  slough.  They  are  of  equal  size.  We  wonder  whether  this  unspotted 
phase  is  a  sexual  color  or  one  due  to  age.  If  this  form  is  a  variant  of  Rana 
pipiens,  is  it  like  albino  or  other  heterochromatic  or  metachroistic  phases  in 
other  groups?  It  is  a  fine  breeding  problem  for  some  Minnesota  student.  (See 
J.  A.  Moore,  Minn.,  1942,  in  "Authorities'  corner.") 

"The  brown  color  changes  of  Rana  burn  si  match  almost  exactly  the  similar 
changes  of  the  Wood  Frog,  Rana  sylvatica,  the  main  difference  being  that 
the  latter  species  seldom  shows  a  strong  green  color  and  always  has  a  black 
patch  at  the  side  of  the  head,  which  is  lacking  in  Rana  burns!'  (A.  C.  Weed, 
Minn.,  1922,  p.  108). 

Color:  Lake  Florida,  Spicer,  Minn.,  Aug.  24,  1930.  Adult.  The  side  of  face, 
top  of  head,  and  down  middle  of  back,  apple  green  or  dull  green-yellow.  On 
the  sides  below  the  costal  folds,  on  rear  of  hind  legs,  courge  green  or  deep  chrys- 
olite green.  Top  of  forearms  and  hind  legs  to  tarsus  mignonette  green.  Groin 
and  rear  of  femur  deep  lichen  green  or  glaucous  green.  Rear  of  tibia  bears  slight 
indistinct  dots  or  lines  of  black  or  light  brownish  olive.  There  are  no  spots  on 
back  except  a  few  minute  inky  dots  between  the  eyes  and  for  an  inch  back- 
ward on  middle  of  back.  Costal  fold,  snout,  and  band  on  upper  jaw  extending 
backward  to  arm  insertion  is  apricot  buff  to  cinnamon-buff,  the  upper  jaw 
tending  toward  mikado  brown  sometimes;  on  the  rear  of  back  between  costal 
folds  are  two  or  three  subfolds  of  orange-citrine  or  citrine.  A  small  black  spot 
on  brachium  near  elbow  is  present  on  one  arm  and  absent  on  the  other.  The 
green  color  of  sides  serves  as  center  of  tympanum  surrounded  by  a  cinnamon- 
brown  circle.  Under  parts  white.  The  eye  is  black  with  outer  circle  of  green 
yellow.  Pupil  rim  baryta  yellow,  broken  below.  Iris  particularly  above  the 
pupil  ring  with  horizontal  band  of  apricot  buff  or  warm  buff.  The  lower 
eye  dotted  with  same  color. 

The  general  color  of  another  frog  is  wood  brown,  the  background  being 
avellaneous  to  light  cinnamon-drab,  with  raised  folds  of  citrine-drab.  The 
costal  folds  are  olive-ocher  to  ecru-olive.  This  frog  has  no  arm  insertion  spots. 

"The  color  is  extremely  variable  in  each  individual  according  to  condi- 
tions of  fear,  etc.,  as  well  as  in  response  to  the  color  of  the  environment  but 
is  very  uniform  in  the  species  as  a  whole.  .  .  .  The  color  of  this  species  is 
usually  some  shade  of  green  or  brown,  varying  from  the  color  of  an  old  water- 
soaked  board  to  a  very  light  mist  gray  and  to  light  apple  green.  About  a  third 


486  HANDBOOK  OF  FROGS  AND  TOADS 

of  the  specimens  have  irregular  black  spots  on  arms  or  legs  or  both.  These 
spots  never  approach  the  condition  of  regular  cross-barring  so  often  seen  in 
Rana  pipiens.  They  look  more  like  blots  of  ink  that  might  have  gotten  there 
accidentally"  (A.  C.  Weed,  Minn.,  1922,  p.  108). 

Structure:  "The  web  of  the  hind  foot  is  quite  variable.  In  the  type  it  appears 
as  a  keel  on  the  sides  of  about  two  and  one  half  joints  of  the  longest  toe,  but  is 
clearly  visible  practically  to  the  end  of  that  toe.  In  some  other  specimens  it  is 
much  larger  and  stretches  almost  directly  across  between  the  tips  of  the  toes. 
There  is  every  degree  of  variation  between  the  two  conditions"  (same,  p.  108). 

Voice:  When  held  just  ahead  of  the  hind  legs,  the  males  inflate  lateral  vocal 
sacs  exactly  like  Rana  pipiens  r  and  do  not  sound  unlike  that  species. 

Breeding:  Eggs,  Tadpoles,  Transformation.  Nothing  recorded. 

Journal  notes:  Aug.  22,  1930.  On  the  third  floor  of  the  Minneapolis  Public 
Library,  Mrs.  Olive  Wiley  has  a  museum  of  reptiles  and  amphibia.  One  puzzle 
to  her  was  a  partly  blue  frog.  It  is  the  green  frog  (Rana  damitans).  In  one 
of  her  aquaria  she  had  a  Rana  she  alluded  to  as  a  wood  frog.  It  had  the  size 
and  general  appearance  of  a  wood  frog,  but  very  manifestly  it  was  the  meadow 
frog  we  sought,  Rana  burnst  of  Weed.  It  is  a  very  bronzy,  beautiful  frog.  Later 
Mr.  O'Connor  showed  me  another  one  which  was  green  on  the  back.  It  too 
had  no  dorsal  spots.  The  first  one  came  from  Coon  Lake  about  10  to  12  miles 
northeast  of  St.  Paul.  This  extends  the  range  of  this  species  outside  the  Minne- 
sota River  drainage. 

Aug.  23.  Arrived  at  Spicer,  Kandiyohi  Co.,  Minn.,  at  5:30  P.M.  Sought  out 
Mr.  M.  F.  Delaske,  who  told  us  that  he  sometimes  had  plain  bait  frogs  with 
no  spots.  Mr.  Oscar  Hillman  did  not  recall  seeing  meadow  frogs  without 
spots. 

Aug.  24.  Started  for  Lake  Florida.  Jens  Larson  had  in  his  bait  box  15  frogs, 
two  half-grown  ones  being  Rana  burnsi.  Started  for  Mr.  Carl  Holm's  place. 
We  stopped  in  a  meadow  where  fringed  gentian,  Lobelia  syphthtica,  smart- 
weed,  Gerardia  pattpercula,  and  other  plants  grew.  Here  took  i  Rana  burn  si 
and  20  Rana  pipiens. 

We  went  down  in  the  meadow  with  Mr.  Carl  Holm.  The  turkeys  were 
there  already.  They  eat  frogs  and  grasshoppers.  We  caught  many  meadow 
frogs.  We  took  in  all  about  5  or  6  Rana  burnsi,  or  about  i  to  every  50  Rana 
pipiens.  Some  of  these  Rana  burnst  are  brown,  others  plain  green.  Mr.  Holm 
said  he  had  found  little  meadow  frogs  without  spots. 

Went  to  South  Florida  sloughs.  Here  we  found  6  or  7  Rana  burnsi.  We 
must  have  seen  200  frogs.  The  ratio  of  Rana  burnsi  to  Rana  pipiens  was  i  to 
28  or  i  to  35.  These  were  in  sedgy  grass.  Most  of  the  frogs  are  doubtless  in 
the  heavier  and  taller  grasses.  Most  of  the  Rana  burnsi  were  half  grown.  We 
took  two  large  males  in  quick  succession.  One  was  uniform  green  and  the 
other  brown.  The  next  day,  when  we  opened  their  can,  they  were  still  of  the 
same  colors. 


FROGS:  RANIDAE  487 

Went  to  Sam  Dilly's  at  Old  Mill  Inn  2  miles  from  New  London.  He  had 
in  his  bait  stock  one  Rana  bnrnsi.  Rana  but  nsi  is  a  beautiful  frog,  brown  or 
green  when  young,  and  equally  variable  when  grown. 

Authorities9  corner: 

A.  C.  Weed,  Minn.,  1922,  p.  108.  L.  Stejneger  and  T.  Barbour,  Check 

A.  C.  Weed,  Minn.,  1930,  p.  43.  List,  1923,  p.  4. 

G.  Swanson,  Minn.,  1933,  p.  154. 

An  embryological  and  genetical  study  of  this  mutant  was  finally  undertaken 
by  J.  A.  Moore,  who  reached  some  interesting  conclusions.  From  his  paper 
(Minn.,  1942,  pp.  408-416),  we  take  the  following  exceipts: 

1.  "A  few  years  ago  a  shipment  of  Rana  ptfnens  arrived  from  the  middle 
west  which  included  several  Rana  bitrnsi.  This  provided  an  opportunity  for 
studying  the  early  development  of  this  non-spotted  variety  and  for  attempting 
to  secure  hybrids  with  typical  Rana  pipiens.  Cross  fertilization  between  bitrnsi 
and  pipiens  was  successful,  and  a  few  tadpoles  were  kept  until  metamorphosis. 
In  the  controls,  btirnsi  9  X  bitrnsi  $ ,  three  tadpoles  transformed.  These  were 
of  two  kinds — one  non-spotted  like  the  parents,  the  other  two  spotted  and  in- 
distinguishable from  ordinary  pipiens.  The  hybrids  of  pipiens  and  bnrnsi 
similarly  gave  rise  to  two  types  of  offspring.  Of  the  24  transforming  in  this 
cross,  13  were  non-spotted  and  n  spotted.  There  was  no  evidence  o{  blended 
inheritance.  Seven  controls,  pipiens  9   X  pipiens  $  ,  differed  in  no  way  from 
their  parents. 

"These  rather  surprising  results  suggested  that  the  non-spotted  frogs  that 
Weed  had  named  Rana  bnrnsi  differed  from  the  typical  Rana  pipiens  by  one 
dominant  gene  carried  in  the  heterozygous  state.  Experiments  with  Rana 
btirnsi  were  resumed  the  following  year.  The  problem  was  assailed  in  two 
ways:  (i)  by  a  study  of  embryonic  temperature  tolerance  and  rate  of  de- 
velopment and  (2)  by  breeding  experiments  to  throw  light  on  the  inheritance 
of  the  pigment  pattern." 

2.  "Morphologically  the  eggs  of  pipiens  and  bnrnsi  are  indistinguishable. 
The  color  and  size  of  the  eggs  as  well  as  the  structure  of  the  jelly  membranes 
are  identical.  Wright  and  Wright  (1933)  have  found  characters  of  this  nature 
useful  in  separating  species  in  many  cases." 

"These  observations  on  egg  size,  type  of  jelly,  rate  of  development,  and 
temperature  tolerance  do  not  reveal  any  difference  between  Rana  pipiens  and 
Rana  bnrnsi.  If  Rana  bnrnsi  proved  to  be  a  cgood  species'  this  would  be  sur- 
prising, but  if  the  difference  between  the  two  is  due  to  a  single  gene  affecting 
pigmentation,  as  was  suggested  by  the  preliminary  experiments,  we  would 
not  necessarily  expect  embryonic  differences  of  the  type  enumerated." 

3.  "Of  the  tadpoles  originally  classed  as  spotted,  90.}  per  cent  gave  rise  to 
pipiens  and  9.7  per  cent  to  bnrnsi.  Of  those  classed  as  non-spotted,  93.7  per 
cent  gave  rise  to  bnrnsi  and  6.3  per  cent  to  pipiens.  The  adult  pigment  dif- 
ferences are  thus  heralded  in  the  larval  stage.  The  exceptions  {bnrnsi  from 


488  HANDBOOK  OF  FROGS  AND  TOADS 

spotted  tadpoles  and  pipiens  from  non-spotted  tadpoles)  can  be  explained  in 
part  by  incorrect  classification  of  very  young  tadpoles." 

4.  "Rana  burnsi  differs  from  Rana  pipiens  by  one  dominant  gene  that  in- 
fluences pigmentation.  Twenty-five  animals  have  been  tested  and  found  to  be 
heterozygous  for  the  mutant  in  question.  Rana  burnsi  should  not  have  the 
status  of  a  species  or  subspecies  but  should  be  reduced  to  synonymy  with 
Rana  pipiens  and  be  referred  to  as  the  'burnsi  mutant!  " 

In  general  we  accept  Moore's  study  even  though  his  burnsi  did  have  a  few 
small  spots  on  the  hind  legs. 

Breckenridge  has  recently  discussed  this  form: 

"Two  color  phases  of  this  species  that  occur  in  Minnesota  differ  so  widely 
from  normal  R.  pipiens  that  they  must  be  mentioned.  These  color  phases  were 
originally  named  as  separate  species  (Weed  1922)  Rana  burnsi  and  Rana 
\andiyohi.  These  species  were  later  reduced  to  subspecies  and  finally  the  terms 
were  dropped  entirely.  Recent  genetic  work  (Moore  1942)  has  proved  that 
the  former  phase  is  a  sport  or  mutant  of  the  typical  form  and  may  be  recog- 
nized simply  as  R.  pipiens,  form  burnsi.  No  similar  work  has  been  attempted 
with  the  other  form  but  since  it  is  probable  that  the  case  is  the  same,  the  form 
is  here  referred  to  as  R.  pipiens,  form  fyandtyohi.  Distinctions  between  the  two 
arc  noted  in  the  section  on  color  phases"  (W.  J.  Breckenridge,  Minn.,  1944, 

P.  82). 

Mottled  Meadow  Frog,  Kandiyohi  Meadow  Frog 

[Phase]  Rana  pipiens  kandiyohi  Weed.  Plate  CHI;  Map  34. 

Range:  "The  type,  number  3066,  Field  museum  of  Natural  History,  was 
received  from  New  London,  Kandiyohi  County,  Minnesota.  Two  other  speci- 
mens, paratypes,  come  from  an  unknown  locality  in  Minnesota.  About  thirty 
other  specimens  were  found  in  lots  of  frogs  from  Rothsay,  Wilkin  County, 
Minnesota,  and  from  Astoria,  Deuel  County,  South  Dakota.  .  .  . 

"The  name  Rana  \andiyohi  is  proposed  with  some  misgivings  because  of 
the  small  knowledge  we  have  of  the  form  and  its  relation  to  other  frogs  and 
to  its  environment.  Localities  from  which  we  have  received  it  are,  with  one 
exception,  in  the  southwest  quarter  of  Minnesota.  The  other  record  is  barely 
across  the  state  line  in  South  Dakota.  It  is  to  be  hoped  that  careful  field  studies 
may  be  made  of  the  frogs  of  the  region  within  a  hundred  miles  of  Big  Stone 
Lake  in  order  that  ecologic  studies  may  supplement  our  present  knowledge 
of  the  group"  (A.  C.  Weed,  Minn.,  1922,  pp.  109-110). 

Habitat:  Open  meadows  and  pastures.  The  young  and  half-grown  frogs 
remain  throughout  the  summer  in  lowlands  or  along  shores  of  lakes  or 
streams. 

Size:  About  same  as  the  common  meadow  frog  (Rana  p.  pipiens). 

General  appearance:  "Typical  specimens  of  this  species  show  a  color  pat- 


FROGS:  RANIDAE  489 

tern  which  suggests  a  blending  of  Rana  pipiens  and  Rana  septentrionalis. 
It  is  as  though  the  black  spots  of  Rana  pipiens  had  been  superposed  on  the 
mottled  color  of  septentrionahs.  The  spots  are  not  as  evenly  rounded  as  in 
pipiens  but  show  a  tendency  to  fuse  with  the  mottlings  between  them.  The 
vermiculate  mottlings  are  carried  down  on  the  legs  and  feet  and  are  there 
combined  with  a  dark  barring  like  that  of  especially  dark  colored  examples  of 
Rana  pipiens. 

"The  mottled  color  of  the  back  is  carried  well  down  on  the  sides  of  this 
species  and  fades  gradually  into  the  white  of  the  under  parts.  The  light  stripes 
along  the  glandular  ridges  are  similar  to  those  in  Rana  pipiens  and  sometimes 
in  Rana  btirnst. 

"The  ground  color  is  represented  by  small  spots  and  lines  between  the 
darker  parts  of  the  mottlings.  It  is  about  the  same  as  in  Rana  pipiens.  Some- 
times the  mottlings  are  as  dark  as  the  spots.  At  other  times  they  are  lighter" 
(A.  C.  Weed,  Minn.,  1922,  p.  109). 

Color:  In  August,  1930,  we  searched  meadows  and  borders  of  lakes  and 
small  streams  from  Spicer,  Minn.,  northward  to  Rainy  Lake  and  westward 
to  Walhalla,  N.D.,  and  then  southward  and  eastward  over  a  different  route 
to  St.  Paul,  Minn.  We  saw  many,  many  meadow  frogs  of  all  sizes  with  a 
wide  variation  in  their  spotting.  Some  had  few  spots,  even  down  to  three  or 
four  small  ones  on  the  back,  others  had  many.  Some  had  long  narrow  spots, 
others  had  round  ones.  In  some  the  spots  were  very  weakly  outlined  so  that 
they  were  much  less  conspicuous.  In  some  there  were  many  dark  specks  and 
dashes  between  the  spots.  In  one  large  frog  the  bars  on  the  tibia  were  entirely 
lacking,  being  replaced  by  many  light  spots  on  a  dark  background.  Along  a 
half-mile  stretch  of  a  canal  at  one  end  of  Green  Lake  at  Spicer,  Minn.,  we 
saw  along  the  shore  200  to  300  small  meadow  frogs.  This  was  in  the  center 
of  Kandiyohi  country.  There  was  wide  variation  in  shape,  size,  number,  and 
arrangement  of  spots,  but  all  appeared  to  us  as  Rana  pipiens. 

Structure:  "The  web  ot  the  hind  foot  is  variable.  In  the  type  it  is  continued 
as  a  rather  broad  keel  to  the  end  ot  the  longest  toe.  In  one  paratype  it  reaches 
about  the  middle  of  the  distal  joint  of  the  longest  toe.  In  the  other  paratype 
it  extends  on  the  next  to  last  joint  of  longest  toe  as  a  very  narrow  keel"  (A.  C. 
Weed,  Minn.,  1922,  p.  109). 

Voice:  "In  conclusion  it  may  be  well  to  mention  that  males  of  Rana  ^andi- 
yohi  kept  alive  in  the  aquarium  have  been  croaking  vigorously  for  some  weeks 
and  that  their  note  is  quite  distinct  from  that  of  some  Rana  pipiens  from  the 
vicinity  of  Chicago.  The  note  of  the  Leopard  Frog  is  more  or  less  a  succession 
of  syllables  and  may  be  almost  represented  by  striking  stones  together  rather 
rapidly.  That  of  Rana  fandiyohi  is  more  in  the  nature  of  a  croak  and  might 
be  represented  by  grinding  two  stones  together  under  considerable  pressure" 
(A.  C.  Weed,  Minn.,  1922,  p.  no). 

Breeding:  Eggs,  Tadpoles,  Transformation.  Nothing  recorded. 


490  HANDBOOK  OF  FROGS  AND  TOADS 

Journal  notes:  Aug.  23,  1930.  Arrived  at  Spicer,  Kandiyohi  Co.,  Minn.,  at 
5:30  P.M.  Sought  out  Mr.  M.  F.  Delaske.  He  told  us  of  Oscar  Hillman  and 
Sam  Dilly  at  Old  Mill  Inn  (2  miles  from  New  London) .  He  told  us  that  frogs 
were  shipped  out  of  Spicer  almost  by  the  ton.  Some  made  $100  in  one  night 
at  frog  catching.  Next  we  went  to  see  Mr.  Hillman.  He  told  us  of  the  immense 
numbers  caught.  He  said  the  mosquitoes  got  thick,  the  farmers  protested, 
and  "the  law  was  put  on  frogs  very  tight."  He  suggested  we  go  to  Carl  Holm's 
meadow  at  Lake  Florida.  More  frogs  were  caught  at  Lake  Florida  than  any 
other  place.  The  land  lies  in  such  a  way  at  Lake  Florida  that  there  is  a  lane 
leading  to  the  lake,  and  here  the  frogs  were  caught  in  great  quantities.  Mr. 
Sam  Dilly  said,  "It  sounds  fishy  but  people  bring  in  frogs  to  Spicer  like  farmers 
drawing  bags  of  grain.  A  man  may  bring  in  8  or  10  such  bags.  They  ship  them 
in  crates  in  tiers."  The  people  in  this  vicinity  shipped  to  F.  J.  Burns  of  Chicago 
and  to  E.  R.  Neuenfeldt  of  Oshkosh.  Many  people  not  only  collected  but 
bought  from  others  to  ship.  Many  Irogs  were  taken  from  Eagle  Lake  and 
Kandiyohi  Lake  near  Willmar. 

August  24.  Started  for  Lake  Florida.  Drove  into  Mr.  Carl  Holm's  place. 
He  gave  us  a  wealth  of  experiences  about  frog  catching  and  frogs.  Mr.  E.  R. 
Neuenfeldt  came  to  his  place  at  first  and  stayed  for  a  time  in  his  home.  At 
first  they  were  paid  3  cents  a  pound  for  frogs,  later  10  cents,  and  once  17  cents. 
It  takes  about  10  to  12  frogs  to  make  a  pound.  Once  he  had  1000  pounds  in 
burlap  bags.  A  freeze  came  and  they  seemed  dead.  He  hated  to  lose  $400.  He 
warmed  some  water  to  tepid  temperature  and  put  it  into  a  barrel.  He  put  one 
bag  of  stiffly  frozen  frogs  into  the  water  and,  lo  and  behold!  they  all  became 
active.  He  thus  revived  his  whole  thousand  pounds. 

When  the  first  frosts  come,  the  frogs  begin  to  leave  the  lowland  meadows 
for  the  lake,  the  young  ones  first,  the  adults  later.  Sometimes  they  delay  so 
long  that  they  get  shut  out  of  the  lake  by  the  formation  of  the  ice.  He  has 
seen  them  moving  in  such  numbers  at  this  season  as  to  form  a  band  two  rods 
wide  and  one-half  mile  long  where  no  one  could  step  without  crushing  frogs. 
They  frequently  put  their  frogs  in  milk  cans.  Sometimes  they  catch  frogs  by 
hand,  sometimes  with  muslin  fences  and  barrel  or  box  traps.  One  night  he 
and  two  other  men  picked  up  750  pounds  by  hand.  It  was  a  cold  night  and 
the  frogs  were  sluggish.  They  took  out  milk  cans  and  could  fill  them  within 
3  inches  of  the  top  without  their  jumping  out.  One  carried  the  can,4two  men 
caught  frogs.  When  catching  with  traps,  they  buy  muslin  and  cut  it  down 
the  middle  and  stretch  it  one  rod  from  the  lake.  The  total  run  may  be  600-700 
yards,  with  a  box  every  hundred  yards.  One  spring  they  had  the  trap  out 
and  caught  turtles,  salamanders,  and  frogs  in  their  boxes.  They  started  col- 
lecting in  1909  and  continued  until  the  collecting  season  was  closed  by  law. 

We  went  down  in  the  meadow  with  Mr.  Carl  Holm.  These  meadows  are 
between  two  ridges  of  hills.  The  meadows  form  a  narrow  strip  or  lane,  a 
capital  place  to  catch  the  frogs  in  fall.  We  examined  many  meadow  frogs.  We 


FROGS:  RANIDAE  49I 

caught  five  or  six  unspotted  ones  and  some  partially  mottled.  One  meadow 
frog  had  tibia  bands  absent  and  tibia  mottled.  Am  not  yet  satisfied  I  have  seen 
a  good  Rana  fyndiyoht.  What  is  Rana  tyndiyohi? 

In  1944  a  live  frog  of  this  phase  came  to  Cornell  with  class  material  from 
Wisconsin.  This  showed  plainly  the  broken  tracery  and  fleckings  between  the 
usual  spots  and  leg  bars. 

Authorities9  corner: 
A.  C.  Weed,  Minn.,  1922,  p.  no. 
A.  C.  Weed,  Minn.,  1923,  p.  28. 
A.  C.  Weed,  Minn.,  1930,  pp.  43-44. 

In  a  letter  of  Oct.  24,  1930,  Dr.  Weed  wrote  to  us:  "Preserved  specimens  of 
^andiyohi  look  much  more  like  pipiens  than  the  individuals  did  in  life.  The 
most  conspicuous  difference  in  the  ones  we  have  here  is  that  the  white  border 
around  the  black  spots  is  more  or  less  definitely  broken  up  into  spots,  which 
are  matched  in  size  and  color  by  similar  spots  elsewhere.  There  seems  also  to 
be  a  definite  tendency  for  black  spots  similar  to  those  of  the  back  to  be  found 
well  down  on  the  flanks  or  even  on  the  sides  of  the  belly.  .  .  . 

"Your  failure  to  find  \andiyohi  at  Florida  Lake  may  be  due  to  its  absolute 
rarity  in  that  region  or  (perhaps)  to  a  difference  in  habitat.  .  .  . 

"In  collections  from  farther  west  the  condition  was  entirely  different.  The 
ratio  of  burnsi  was  about  the  same  but  fyandiyohi  tended  to  equal  pipiens  in 
numbers  and,  as  I  remember  it,  actually  exceeded  pipiens  in  one  lot  of  several 
thousand  from  South  Dakota.  .  .  . 

"In  thinking  over  my  experiences  in  collecting  these  frogs,  it  comes  definitely 
in  my  mind  that  if  I  were  looking  for  a  large  stock  of  \andiyohi  I  should  start 
somewhere  in  the  vicinity  of  Lake  Traverse  and  Big  Stone  Lake  and  work 
westward  toward  the  region  where  it  becomes  too  dry  for  frogs  to  live.  I  should 
expect  to  find  a  region  where  pipiens  is  about  as  rare  as  J^andiyohi  is  in  some 
parts  of  Minnesota." 

Since  this  letter  only  Swanson's  (193?)  and  Breckenridge's  (1944)  contribu- 
tions are  pertinent. 

"The  vermiculated  form  (\andiyohi)  Weed  reports  from  Kandiyohi  and 
Wilkin  counties.  It  has  not  been  noted  in  the  Minneapolis  region"  (G.  Swan- 
son,  Minn.,  1935,  p.  154). 

"The  dark  color  phase  of  the  leopard  frog,  form  \andiyohi,  is  characterized 
by  a  marked  darkening  of  the  interspaces  between  the  black  spots  of  normal 
R.  pipiens.  On  the  hind  legs,  the  black  spots  arc  often  much  broken  with  coarse 
fleckings  of  black  and  greenish  or  yellowish.  The  result  is  a  very  dark  frog, 
with  the  black  spots  of  the  normal  pattern  indistinctly  outlined  in  broken  light 
lines.  The  dorso-lateral  ridges  are  often  metallic  in  appearance.  The  few  rec- 
ords of  this  dark,  mottled  form  of  the  leopard  frog  are  rather  widely  scattered 
over  the  prairie  section  of  Minnesota,  with  one  isolated  record  in  Hennepin 
County  in  the  eastern  part  of  the  state.  ...  A  fisherman's  frog  box  containing 


492  HANDBOOK  OF  FROGS  AND  TOADS 

250  frogs  was  examined  near  Granite  Falls  and  was  found  to  contain  five  or 
2  per  cent  of  this  dark  form.  On  the  basis  of  present  data  the  author  considers 
form  tyndiyohi  to  be  much  less  common  than  form  burnsi  and  more  restricted 
in  distribution"  (W.  J.  Breckenridge,  Minn.,  1944,  p.  84). 

Southern  Meadow  Frog,  Southern  Leopard  Frog,  Spring  Frog,  Spotted  Frog, 

Water  Frog,  Shad  Frog 

Rana  pipiens  sphenocephala  (Cope).  Plate  CIV;  Map  34. 

Range:  Southeastern  states,  north  along  the  coast  to  Virginia  or  New  Jersey; 
up  Mississippi  River  to  Indiana  and  Kentucky. 

Habitat:  Ponds,  runs,  canals,  river  swamps,  and  overflowed  roads  and 
ditches.  In  a  large  swamp  it  is  ideally  on  the  edge  of  cypress  ponds  and  bays 
or  small  pools  at  an  island's  edge. 

"Margins  of  streams  and  lakes,  and  marshy  spots,  common"  (O.  C, 
Van  Hyning,  Fla.,  1933,  P-  4)- 

"When  alarmed  they  usually  make  six  or  eight  very  long  leaps,  pivoting 
at  each  landing.  Among  thick  palmettos  I  have  seen  them  cover  some  distance 
in  their  precipitous  retreats  without  ever  touching  the  ground,  each  jump  be- 
ginning and  ending  on  one  of  the  broad  leaves.  Rarely  one  dives  into  the  water 
when  cornered,  usually,  however,  reappearing  at  the  water's  edge  and  con- 
tinuing the  retreat  on  land"  (A.  F.  Carr,  Jr.,  Fla.,  1940^  p.  67). 

Size:  Adults,  2-3  ]/4  inches.  Males,  49-78  mm.  Females,  53-82  mm. 

General  appearance:  It  is  like  the  meadow  frog  R.  p.  pipiens,  but  usually 
with  a  clear-cut,  distinct  white  spot  in  the  middle  of  the  eardrum.  It  is  an  alert, 
active,  long-legged,  and  long-snouted  spotted  frog. 

It  is  usually  glistening  white  below,  but  on  the  Florida  Keys  a  form  occurs 
that  frequently  has  bronzy  throat  and  clouded  or  mottled  belly;  and  young 
there  may  have  spotted  throats  and  under  parts.  (See  R.  pipiens  complex,  p. 
502.) 

Color:  Male.  Okefinokee  Swamp,  Ga.,  July  3, 1921.  Olive,  jade,  or  grass  green 
on  back;  pale  chalcedony  yellow  on  upper  lip;  pale  dull  green-yellow  costal 
fold;  center  of  tympanum  chalcedony  yellow.  Some  amber  yellow  or  primulinc 
yellow  on  underside  of  forearm,  little  on  side  of  groin,  on  outer  side  of  hind 
legs,  and  back  of  tarsus.  Under  parts  white.  Thumb  with  black  swelling.  Iris, 
back  part  chartreuse  yellow;  upper  part  vmaceous-buff;  inner  ring  amber 
yellow  or  wax  yellow;  rest  black. 

Female.  Okefinokee  Swamp,  Ga.,  June  21, 1921.  Top  of  head  to  back  of  eye, 
cinnamon,  Prout's,  or  snuff  brown  to  bister,  sudan,  or  brussels  brown.  Same 
color  along  inner  and  outer  sides  of  costal  fold,  on  upper  jaw  from  snout  to 
front  of  eye,  on  upper  fore  limb,  and  on  ground  color  of  back,  at  times  be- 
coming the  predominant  ground  color.  Background  of  back  dark  greenish 
olive  to  ivy  green.  Spots  of  back,  sides,  arms,  and  legs  black.  Those  of  back  with 


FROGS:  RANIDAE 


493 


Plate   CHI.  Rana  ptpiens   (tyndiyohi) 
(X%)-  Females,  from  Wisconsin. 


Plate  CIV.  Rana  pipiens  sphenocephala. 
i.  Male  (X%).  2.  Female  (X%).  3-  Fe- 
male (x%). 


494  HANDBOOK  OF  FROGS  AND  TOADS 

a  thin  light  green-yellow  border.  Under  parts  white.  Throat  and  pectoral 
region  dulled  with  blackish.  Lower  jaw  rim  marguerite  yellow  or  seafoam 
yellow  with  black.  Same  color  for  upper  jaw  below  the  stripe  from  lower  eye 
to  arm  insertion.  Above  the  sudan  brown  of  the  side  of  snout  and  in  front  of 
eye  through  nostril  is  a  blackish  or  greenish  color.  Two  or  three  marguerite 
or  seafoam  yellow  spots  behind  the  eye.  Stripe  from  under  eye  over  angle  of 
mouth  and  just  beneath  tympanum  to  over  arm  insertion  marguerite  yellow 
or  seafoam  yellow.  Tympanum  argus  brown  with  marguerite  yellow  or  sea- 
foam  yellow  center.  Iris  in  front,  below,  and  behind  pupil  black  with  some 
pale  vinaceous  lilac  or  light  pinkish  lilac;  rim  pinard  yellow;  iris  above  pupil 
maize  yellow;  very  top  of  eye  with  a  small  black  area. 

Structure:  Head  longer  in  proportion  to  body  than  in  R.  piptens;  snout 
acuminate  or  pointed;  fourth  toe  shorter  than  in  R.  pipiens  and  more  regularly 
placed;  fewer  spots  and  more  vermiculations  on  sides  of  R.  p.  sphenocephala; 
dusky  fleckings  on  the  under  parts  more  common  in  R.  p.  sphenocephala; 
lower  lip  with  dark  spots;  vocal  sacs  on  either  side  between  arm  and  tympa- 
num. 

Voice:  The  call  is  three,  four,  or  five  guttural  croaks  with  two  or  three  clucks 
afterwards.  The  process  may  occupy  5  or  6  seconds.  They  are  shy,  croak  at 
night,  and  even  a  big  chorus  is  quickly  stopped  as  one  approaches  the  pond. 
They  may  croak  from  the  surface,  beneath  the  surface,  or  from  perches  on 
logs,  sticks,  or  around  the  bases  of  bushes.  The  head  and  upper  back  are  usually 
emergent.  The  croak  itself  comes  from  the  swift  inflation  and  deflation  of  the 
vocal  sacs,  which  are  on  either  side  between  forelegs  and  tympanum.  At  n  :2o 
P.M.,  May  16, 1921,  we  heard  a  Rana  p.  sphenocephala  chorus.  Strange  we  did 
not  hear  them  when  we  were  near  the  pond  but  distance  had  to  sift  out  the 
calls.  We  had  less  success  in  photographing  croaking  males  than  with  any 
other  species  encountered.  July  13,  1922,  at  12:30-1:00  A.M.,  we  awoke  and 
heard  a  great  chorus  northwest  of  our  spring.  Here  in  grassy  borders  of  open 
spots  were  plenty  of  meadow  frogs.  In  the  same  place  Aug.  9,  heard  two  im- 
mense choruses,  one  at  5:00  A.M.,  and  another  at  4-5  A.M.  Went  to  the  latter 
congress.  Rarely,  they  are  heard  in  the  daytime  before  or  after  a  storm  passes. 

Breeding:  This  species  breeds  from  February  to  December,  the  crest  coming 
from  April  to  August.  The  egg  mass  is  a  plinth,  5-6  inches  (125-150  mm.) 
wide,  1-2  inches  (25-50  mm.)  deep,  attached  to  stems  of  plants  or  to  sticks, 
submerged.  The  egg  is  Vi(J  inch  (1.6  mm.),  the  envelopes,  %  inch  (3.2  mm.) 
and  %  inch  (5.4  mm.).  The  tadpole  is  large,  3  inches  (74  mm.),  the  tail  with 
conspicuous  black  blotches  as  transformation  approaches.  The  tooth  ridges  are 
%  or  %.  After  67  to  86  days,  the  tadpoles  transform  from  April  to  October 
at  %-i%  inches  (20-33  mm.). 

"They  may  breed  every  month  of  the  year.  The  eggs  are  usually  laid  in 
masses  of  vegetation  (algae,  Websteria,  Persicaria,  Hypericum,  etc.)  at  the 


PROGS:  RANIDAfi 

water's  edge  or  in  the  shallower  portions  of  ponds  and  ditches"  (A.  F.  Carr,  Jr., 
Fla.,  1940,  p.  68). 

"It  is  an  early  breeder,  laying  its  eggs  at  Raleigh  in  February  and  March, 
usually  starting  a  little  later  than  the  chorus  frogs,  but  often  the  two  are  singing 
at  the  same  time  as  their  breeding  largely  coincides.  However,  the  eggs  arc 
occasionally  laid  in  late  autumn"  (C.  S.  Brimley,  N.C.,  1940,  p.  27). 

Journal  notes:  In  1921  we  made  these  notes  on  places  of  ovulation.  On  April 
24,  we  found  them  in  a  cypress  pond.  Center  of  pond  is  clear  (alligator  hole) ; 
then  came  a  circle  of  Pontederia  in  which  are  toad  and  Rana  p.  sphenocephala 
eggs;  a  circle  of  sedges;  next  came  bushes;  a  thin  line  of  gums;  and  finally 
the  pines.  On  April  29,  Noah  took  me  to  where  he  had  found  frogs'  eggs.  They 
were  about  8  feet  from  the  edge  of  the  pond  in  water  4-6  inches  deep.  This 
mass  just  hatching  Noah  found  ySsterday  morning.  Two  more  isolated  masses 
found,  each  at  or  just  below  the  surface  and  encircling  the  li/ards'  tail 
(Saiirurus)  which  is  now  in  bloom.  Water  on  the  surface  90°. 

On  June  i  we  remarked  that  Rana  p.  sphenocephala  were  hopping  around 
the  island  after  breeding,  also  that  they  were  becoming  browner  when  afield 
and  as  the  season  progressed. 

Sept.  1 1, 1929.  About  dusk  stopped  near  Lyons,  Ind.,  near  home  of  Gless  J. 
Deckard.  Heard  many  meadow  frogs  in  a  pond  back  of  his  place.  Found 
many  of  them  at  edge  of  the  pond  in  grass  or  in  little  depressions. 
At  times,  though  not  croaking,  they  kepi  sac  inflated.  They  have  more 
pointed  snouts  than  our  northern  meadow  frog  and  all  have  light  centers  to 
the  tympanums.  Is  there  another  breeding  period  in  the  fall?  Saw  no  females. 
Do  supernumerary  males  or  young  males  approaching  breeding  si7,e  croak 
the  fall  before,  like  young  roosters  learning  to  crow?  One  answer  came  the 
very  next  day.  Went  on  to  Olney.  Drove  on  the  "slab  road,"  about  a  mile  south 
of  the  town  lights.  Here  found  a  pond  beside  the  road  and,  lo  and  behold! 
there  were  about  ten  egg  complements  of  meadow  frogs. 

Authorities9  corner: 
W.  Bartram,  Gen.,  1791,  pp.  278-279. 
J.  D.  Kilby,  Fla.,  1945,  pp.  79-80. 

Virginia.  (M.  K.  Brady,  1927,  p.  27.)  "Leopard  frogs  in  this  locality  |  Dismal 
Swamp  |  have  all  the  characteristics  of  sphenocephala.  They  suggest  ptpiens 
even  less  than  the  leopard  frogs  of  the  Potomac  drainage  do." 

North  Carolina.  (C.  S.  Brimley,  1940,  p.  27.)  "Nearly  allied  to  this  is  the 
Northern  Leopard  Frog  (Rana  pipiens)  which  grows  to  a  larger  size  (maxi- 
mum length  head  and  body  about  four  inches  instead  of  about  three  in  south- 
ern form),  usually  has  a  blunter  snout,  and  generally  does  not  have  a  white 
spot  on  the  tympanum." 

South  Carolina.  (E.  B.  Chamberlain,  19^9,  p.  32.)  "Although  Holbrook  did 
not  distinguish  this  southeastern  form  from  the  more  northerly  pipiens  his 


496  HANDBOOK  OF  FROGS  AND  TOADS 

plates  of  'Rana  halecina  show  the  light  spot  at  the  center  of  the  tympanum, 
usually  considered  as  diagnostic  of  sphenocephala.  An  occasional  specimen 
from  the  Upper  Piedmont  seems  to  show  an  approach  to  pipiens,  but  I  have 
seen  no  typical  example  of  the  latter  form  from  this  state.  .  .  ." 

Georgia.  (A.  H.  Wright,  1931,  p.  434.)  "This  form  seems  a  Lower  Austral 
offshoot  of  Rana  pipiens  extending  from  New  Jersey  to  Texas  and  up  the 
Mississippi  to  Indiana."  (Still  our  opinion  after  14  years.) 

Florida.  (S.  Springer,  1938,  p.  49.)  "In  a  recent  paper  on  the  status  of  the 
leopard  frogs  (Herpetologica  i,  1937:  84-87)  Mr.  Carl  F.  Kauffeld  says  that 
Rana  brachycephala  attains  a  greater  size  than  either  pipiens  or  sphenocephala. 
Over  most  of  its  range  I  have  no  doubt  that  sphenocephala  is  relatively  small. 
In  the  area  west  of  Myakka  River  on  the  west  coast  of  Florida,  however, 
sphenocephala  attains  a  very  large  size.  Specimens  having  a  body  length  of  four 
and  one-half  inches,  measured  from  snout  to  anus,  are  quite  common,  and  I 
have  seen  a  few  specimens  with  a  body  length  of  five  inches.  One  of  these 
weighed  half  a  pound." 

Mississippi.  (M.  J.  Allen,  1932,  p.  10.)  "An  examination  of  several  hundred 
frogs  leads  to  a  conclusion  that  the  majority  of  specimens  agree  with  descrip- 
tions of  R.  sphenocephala  with  a  few  exhibiting  tendencies  toward  intergrada- 
tion,  and  a  small  percentage  having  some  of  the  characters  of  Rana  pipiens" 

Louisiana.  (Viosca  has  consistently  called  his  local  torms  different  from 
northeast  R.  ptpiens.) 

Oklahoma.  (A.  N.  Bragg,  19423,  p.  18.)  "Some  call  this  frog  Rana  pipiens 
Schreber;  others  believe  both  species  occur  in  Oklahoma.  It  is  the  author's 
opinion  that  R.  sphenocephala  is  the  common  leopard  frog  of  the  state  but 
that  some  specimens  in  the  northwestern  part  may  be  pipiens.  The  latter  is  a 
larger,  broader-headed  species." 

Illinois.  (D.  W.  Owens,  1941,  p.  184.)  "In  order  to  test  the  relations  of  the 
Macoupin  County  specimens  to  the  leopard  frog  of  the  Chicago  Region  and  to 
specimens  recorded  as  Rana  sphenocephala  from  southern  Illinois,  I  have 
measured  the  length  of  body  and  the  tibia  in  all  of  the  Illinois  specimens 
available  in  Field  Museum.  .  .  .  The  tibia  and  body  measurements  expressed 
as  the  t/b  ratio  are  as  follows : 

No.  of  Extremes  Average 

specimens 

Chicago  Region — pipiens  63  .5o-.6i  .55 

Southern  Illinois — sphenocephala  53  .^-.62  .58 

Macoupin  County  specimens  10  -54--63  -59 

I  concluded  that  the  Macoupin  County  leopard  frog  should  be  referred  to 
sphenocephala,  the  longer-legged  and  more  sharp-snouted  form  of  southeastern 
United  States.  .  .  ." 


FROGS:  RANIDAE 


497 


Indiana.  (F.  N.  Blanchard,  1925,  p.  372.)  "All  the  leopard  frogs  collected  are 
referred  to  this  species  rather  than  to  R.  ptpiens  because  they  look  more  like  the 
southern  than  the  northern  leopard  frogs.  They  are  very  different  in  appear- 
ance from  the  leopard  frogs  of  Michigan.  The  spots  on  the  back  are  very  small 
and  are  without  light  outlines.  The  head  averages  a  little  longer  than  on 
leopard  frogs  from  Michigan,  its  length  being  contained  in  the  total  length 
two  and  one-half  to  three  times.  The  probability  is  that  R.  sphenocephala  inter- 
grades  with  R.  pipiens  somewhere  north  of  Southern  Indiana."  (See  also  Swan- 
son,  Ind.,  1939,  pp.  688-689.) 

Kentucky.  (  W.  A.  Welter,  K.  Carr,  R.  W.  Harbour.)  We  have  seen  some  of 
the  material  these  workers  identify  as  Rana  p.  sphenocephala. 

In  1944  J.  A.  Moore  (Gen.,  1944,  pp.  549-369)  gave  us  a  scholarly  study  of 
geographic  variation  in  Rana  pipiens  of  eastern  North  America.  Some  com- 
ments on  Dr.  Moore's  article  follow : 

1.  When  Miss  Dickcrson  (1906)  of  the  American  Museum  featured  Rana 
sphenocephala,  there  were  young  skeptics.  Never  have  we  been  strong  for  it 
as  a  separate  species.  Probably  R.  p.  sphenocephala  is  a  better  handle.  But  in 
spite  of  our  (1906)  skepticism,  we  soon  began  to  recognize  a  southern  leopard 
frog.  In  our  40  years'  travels  in  the  United  States  we  have  seen  these  frogs  in 
their  places,  and  we  still  recognize  sphenocephala,  call  it  species,  subspecies,  or 
race. 

2.  Sphenocephala  sorts  out  the  best  of  all  the  pipiens  complex.  Before  we 
read  Dr.  Moore's  paper  or  legends  we  sorted  his  ptptens  and  sphenocephala 
illustrations.  Strange  to  say,  we  were  over  go  per  cent  right  on  sphenocephala 
whether  the  frogs  were  facing,  straight  lateral,  dorsal,  or  inclined  toward  or 
away  from  the  photographer. 

3.  Much  of  Dr.  Moore's  tilting  is  with  Kauffeld's  establishment  of  three 
races  along  our  Atlantic  coast.  We  never  indorsed  that  solution. 

4.  At  times  we  have  questioned  Cope's  use  of  external  vocal  sacs  in  separa- 
tion of  R.  pipiens  forms.  It  is  strange  that  Moore  finds  that  "from  southern 
New  York  and  south  along  the  Atlantic  coast  the  males  possess  well-developed 
external  vocal  sacs.  ...  In  the  Mississippi  Valley   males  from   southern 
Indiana,  Oklahoma,  Arkansas  and  Mississippi  have  well  developed  external 
vocal  sacs.  .  .  ." 

This  reads  much  like  our  range  characterization  for  Rana  p.  sphenocephala. 

5.  "Males  from  southern  New  York  to  northern  Florida  lack  oviducts,  .  .  . 
The  same  is  true  for  males  in  Louisiana,  Mississippi,  eastern  Texas  and  in  the 
Mississippi  Valley  north  to  southern  Indiana,  Illinois  and  Missouri"  (J.  A. 
Moore,  p.  562).  Compare  Moore's  map  of  males  without  oviducts  with  our 
range  map.  One  could  substitute  his  map  for  ours  if  one  eliminated  southern 
peninsular  Florida. 

6.  These  characters  (morphological  and  physiological)  cannot  be  read  out 


498 


HANDBOOK  OF  FROGS  AND  TOADS 


of  the  picture.  Still  we  call  R.  p.  sphenocephala  separate  because  of  the  "look 
of  the  beast."  To  return  to  these  vocal-sac  and  without-oviduct  males  of 
JR.  p.  sphenocephala:  Could  it  be  that  sphenocephala— 

(a)  Matures  smaller? 

(b)  Matures  faster  because  it  can  grow  the  whole  season,  R.  p.  pipicns  does 
not  and  cannot  because  of  northern  winters? 

(c)  Breeds  every  month  in  the  year,  is  a  more  active  breeder? 


Rana  pipicns  complex 
Plates  CV-CXV;  Map  34. 

A  comparison  of  Map  34,  showing  the  hypothetical  distribution  of  various 
Rana  forms,  with  a  geological  map  of  the  United  States  suggests  that  Rana 
piptens  frogs  may  find  reasons  for  changing  their  coats  in  the  chemical  and 
structural  changes  of  their  habitats  in  the  different  sections  of  the  country. 


Map  33 


a 


500  HANDBOOK  OF  FROGS  AND  TOADS 

Is  pipiens  one  form,  as  Kellogg,  Bogert,  Moore,  and  others  believe,  or  many 
subspecies?  For  many  years,  we  were  of  the  first  school.  Today,  we  are  in 
balance,  in  suspended  judgment.  We  feel  that  R.  p.  sphenocephala  is  as  good 
a  form  as  many  forms  in  snakes,  better  than  some  in  birds  and  mammals, 
and  as  good  as  some  drainage  forms  in  fishes.  In  1925,  we  felt  R.  fisheri  prob- 
ably a  good  form,  but  at  present,  are  the  most  striking  specimens  of  this 
limited  population  gone?  We  present  this  rambling  treatment  to  provoke 
thorough  study — study  of  all  the  collections  plus  travel  experience  with  these 
very  forms,  their  eggs,  tadpoles,  and  life  histories.  We  are  past  appraisals  of 
alcoholic  specimens  alone.  Those  began  75  years  ago. 

We  hold  the  problems  to  be  the  isolated  populations  in  the  West  and  agree 
with  Schmidt  (Schmidt  and  Smith,  Tex.,  1944,  p.  81),  "The  senior  author  is 
convinced  that  Rana  sphenocephala  can  be  distinguished,  and  that  the  essential 
problem  lies  in  the  Trans-Mississippian  populations,  in  which  characters  dis- 
tinctive of  pipiens  and  sphenocephala  in  eastern  North  America  appear  in 
various  mixtures." 

A.  The  color  phases  or  mutant  forms,  Plates  CV;  CVI;  CVII. 

1.  Bttrnsi  mutant.  (See  pages  483-488  for  an  account  of  this  color  phase.)  We 
never  considered  this  mutant  and  kandtyoht  valid  species  or  subspecies,  but  we 
included  them  in  earlier  editions  of  the  Handbook^  to  provoke  some  mid- 
western  geneticist  to  drop  Drowphtla  and  work  on  an  immediate  backyard 
form,  but  it  remained  for  someone  in  New  York  City  to  make  the  study. 
Many  an  American  herpetologist  on  one  specimen  or  a  few  specimens  has 
made  new  species  on  as  slender  grounds  as  our  old  classmate,  Weed. 

In  elementary  class  lots  of  many  hundreds  of  frogs  from  Vermont  and  other 
parts  we  have  occasionally  seen  unspotted  R.  pipiens,  just  as  individual  evittate 
Hyla  cinerea  appear  in  each  southern  state. 

2.  Kandiyohi  phase.  (See  pp.  488-492  for  an  account  of  this  form.) 

B.  Generally  accepted  form,  R.  p.  sphenocephda.  (See  pages  492-498  for  an 
account  of  this  subspecies.) 

Most  herpetologists  since  Miss  Dickerson  have  accepted  this  southeastern 
meadow  frog  as  distinct  and  call  it  a  race,  variety,  subspecies,  or  species. 

It  is  true  that  Boulenger  has  written:  "I  am  therefore  unable  to  divide  the 
species  in  minor  groups  with  any  precision,  and  must  leave  the  matter  in 
abeyance  for  the  present."  But  he  concluded  his  R.  halecina  account  as  follows: 
"A  precise  diagnosis  of  the  var.  sphenocephala  is  still  a  desideratum,  as  Miss 
Dickerson's  definition  .  .  .  seems  to  me  insufficient,  in  view  of  the  variation 
in  specimens  referable  to  the  typical  form"  (G.  A.  Boulenger,  Gen.,  1920,  p. 
439) .  KauflEeld  accepts  this  form  as  a  species  but  he  would  put  the  Piedmont 
specimens  from  "extreme  southeastern  New  York,  Long  Island,  southern 
Connecticut,  New  Jersey  except  the  northwestern  portion,  southeastern 
Pennsylvania,  Delaware,  Maryland,  south  throughout  the  Coastal  Plain  and 
west  into  Texas"  as  Rana  pipiens  Schreber  and  the  forms  from  the  Fall  line 


CF.  (X%)-  From  lake  Florida,  Minn, 


Plate  CVI.  pipiens  (burnsi).  (X%)*  From  Lake  Florida,  Minn. 


Plate  CVU.  Left.  Rana  pipiens  (\andiyohi).  From  Wisconsin.  Right.  Rana 
pipiens  (fyndiyohi)  intermediate.  From  Lake  Florida,  Minn. 


502  HANDBOOK  OF  FROGS  AND  TOADS 

to  the  coast  as  R.  sphenocephala.  We  are  not  of  his  school  of  interpretation,  yet 
we  thank  him  for  his  compact  paper,  which  bravely  touched  off  a  sudden  spurt 
of  activity  in  R.  pipiens  study.  In  the  same  way,  we  admire  the  thoroughness 
of  Moore's  paper  yet  wonder  whether  some  authors  have  had  extensive  field 
experience  throughout  the  range  of  R.  p.  sphenocephala.  For  years  we  have 
been  inebriated  from  alcoholic  inspirations  (the  same  thoughts  some  of  the 
old  masters  had).  Get  out  in  the  field  of  the  species  treated. 

When  Moore  disagrees  with  Mittleman  and  Gier  regarding  their  R.  her- 
landieri  (and  we  agree  with  Moore  on  this  point),  his  clincher  is :  "In  fact  there 
are  greater  differences  between  species  from  Oklahoma  and  part  of  Texas  than 
between  the  former  and  specimens  from  southern  Indiana  and  parts  of  New 
Jersey."  Of  course  he  destroys  R.  p.  sphenocephala,  but  we  smile  to  recall  that 
Oklahoma,  southern  Indiana,  and  New  Jersey  are  within  the  range  of  jR.  p. 
sphenocephala ,  as  is  Lake  Caddo  (on  the  boundary  of  Texas  and  Louisiana) 
and  possibly  Belton,  Tex.  We  have  traveled  over  these  areas  several  times.  Be- 
fore one  discusses  Elaphe,  Crotaltts,  or  the  range  of  any  form,  well-recognized 
or  debatable,  one  should  travel  over  the  area.  Because  we  have  rarely  seen 
an  unspotted  R.  p.  pipiens  from  Vermont  or  Texas,  we  would  not  assert  that 
a  regular  mutant  population  (burnsi)  did  not  exist  in  Minnesota,  without 
visiting  the  region  in  question.  We  need  more  of  this  type  of  investigation  to 
reinforce  the  cloister  of  university  and  museum. 

We  append  A.  N.  Bragg's  (N.  Mex.,  1941,  p.  116)  trenchant  remark:  "After 
observing  the  two  forms  in  the  field,  I  feel  certain  that  the  frog  about  Las 
Vegas,  New  Mexico  is  very  similar  to,  if  not  identical  with,  the  grass-frog  of 
New  England  (Rana  brachycephala)  of  the  recent  checklist  and  the  leopard 
frog  of  Wisconsin,  and  different  from  that  of  Oklahoma  which  I  call  Rana 
sphenocephala  (Cope)." 

C.  Eastern  puzzles, 
i.  Florida  Keys. 

On  March  20  and  21, 1934,  we  made  the  following  journal  notes:  Last  night 
went  along  the  railroad  ditches.  Saw  R.  p.  sphenocephala  on  way  to  Key  West. 
Looked  in  sinkholes,  i.e.,  honeycombed  rock,  containing  water  and  in  them 
were  meadow  frogs.  In  a  shallow  covered  well  found  several  in  holes  along  the 
sides.  On  a  Key  near  Big  Pine  Key  were  numberless  R.  p.  sphenocephala.  The 
adults  were  bronzy  on  throat  with  belly  much  clouded  and  the  entire  under 
parts  as  cloudy  as  a  Heckscher's  frog  or  a  young  R.  clamitans.  The  half-grown 
ones  have  very  spotted  throats  and  under  parts.  Someday  someone  with  as  close 
discrimination  of  details  as  now  obtains  in  birds  or  snakes  (scales  to  count) 
may  make  a  new  subspecies  of  this  colored  phase.  We  put  the  frogs  in  our 
covered  fish  can.  Later  when  we  photographed  them  we  wrote  in  the  photo- 
graphic record:  "The  venter  has  already  grown  lighter  but  legs  are  still 
mottled.  Big  Pine  Key."  In  a  day  or  so  some  died.  These  looked  almost  exactly 
like  ordinary  R.  p.  sphenocephala. 


FROGS:  RANIDAE  503 

2.  Common  northern  form  (pipiens). 

In  1942  Moore  (Gen.,  19423,  VI,  201)  reviewed  the  northeastern  situation 
as  follows:  "Unfortunately  taxonomists  are  not  in  agreement  on  the  status  of 
these  frogs.  Some  years  ago  Cope  (1889)  divided  Rana  virescens  (an  old  name 
for  Rana  pipiens)  into  three  subspecies,  R.  v.  brachycephala,  R.  v.  pipiens  and 
R.  v.  sphenocephala.  Later  workers  for  the  most  part  did  not  recognize  the 
subspecies  brachycephala  and  called  all  individuals  from  the  northern  states 
and  Canada  Rana  pipiens.  The  southern  variety  on  the  other  hand  was  raised 
to  specific  rank,  Rana  sphenocephala.  The  differences  between  these  two  'spe- 
cies' were  so  slight  that  many  competent  herpetologists  would  have  difficulty 
in  identifying  a  given  specimen  unless  the  locality  was  plainly  indicated  on  the 
label.  Kellogg  (1932)  examined  an  extensive  series  of  frogs  from  many  parts 
of  the  country  and  thought  it  best  to  lump  them  as  Rana  pipiens.  Shortly 
after  this  Kauffeld  (1937)  suggested  that  all  three  of  Cope's  subspecies  be 
raised  to  full  specific  rank  —  Rana  brachvccphala,  Rana  pipiens  and  Rana 
sphenocephala.  The  diagnostic  characters  given  by  Kauffeld  have  not  been 
applied  successfully  by  all  taxonomists  (Trapido  and  Clausen  1958;  Grant 


C.  F.  Kauffeld  certainly  deserves  special  thanks  for  his  two  papers  in 
Herpetologica  (Gen.,  1936,  p.  n;  1937,  pp.  84-87).  He  started  a  lot  of  writing 
if  nothing  more.  He  wrote: 

"For  some  inexplicable  reason,  Rana  brachycephala  of  Cope  never  gained 
the  recognition  it  deserved  although  sphenocephala  of  the  same  author,  a  form 
much  less  distinct  from  true  pipiens  than  brachycephala,  has  been  universally 
accepted.  Rana  brachycephala  remained  in  complete  obscurity  from  the  time 
of  its  description  (Cope  1889,  pp.  403-406).  No  one,  so  far  as  I  can  find,  even 
bothered  to  throw  it  into  synonymy  until  Kellogg  (1929,  p.  20  })  placed 
brachycephala  as  well  as  sphenocephala  in  the  synonymy  of  pipiens.  There  is 
no  justification  for  treating  either  of  these  two  forms  in  this  way  and  brachy- 
cephala must  be  revived  to  supply  a  name  for  the  leopard  frog  which  has  so 
often  been  erroneously  figured  and  described  as  pipiens  (Dickerson  1908,  pi. 
ii  ;  Wright  and  Wright  1933,  pi.  71)." 

Schmidt  does  not  endorse  the  use  of  R.  brachycephala  for  our  northeastern 
meadow  frog.  "Kauffeld's  remarks  (1937,  P-  84)  about  Rana  attstricola, 
R.  pipiens  burnsi,  and  R.  \andiyohi,  and  his  choice  of  brachycephala  as  the 
name  of  the  northeastern  'pipiens'  are  unintelligible  to  us"  (K.  P.  Schmidt  and 
T.  F.  Smith,  Tex.,  1944,  p.  81). 

To  quote  Kauffeld  again  before  we  proceed  to  the  discussion:  "As  the  ma- 
terial which  Cope  had  at  the  time  he  described  brachycephala  (which  by  the 
way,  numbered  182  specimens  and  not  15  Kellogg  loc.  cit.  mistakenly  esti- 
mated) [Dr.  Kellogg  clearly  says  2  adults,  i  young,  and  12  tadpoles  in  the  type 
lot  no.  3363.—  A.  H.  W.]  this  species  occurs  in  the  Transition  Zone,  partic- 
ularly the  Alleghanian  Division,  the  southern  extension  of  the  Canadian,  and 


504  HANDBOOK  OF  FROGS  AND  TOADS 

the  Upper  Austral  west  of  the  Appalachian  Highlands.  Other  material  sub- 
stantiates this  distribution.  .  .  .  Schreber's  type  of  pipiens  was  undoubtedly 
from  the  Carolinian  of  southern  New  York.  We  may  define  the  range  of 
brachycephala  and  piptens  as  follows:  Rana  brachycephala  (Cope),  Southern 
Canada  and  New  England  except  extreme  southern  Connecticut,  New  York 
except  the  southeastern  portion,  northwestern  New  Jersey,  northern  and  west- 
ern Pennsylvania,  West  to  the  Pacific  Coast  States.  Rana  piptens  Schreber, 
Extreme  southeastern  New  York,  Long  Island,  southern  Connecticut,  New 
Jersey  except  the  northwestern  portion,  southeastern  Pennsylvania,  Delaware 
and  Maryland,  south  throughout  the  coastal  Plain  and  west  into  Texas" 
(Kauffeld,  Gen.,  1936,  p.  n;  1937,  pp.  84-87).  [Neither  his  description  nor 
his  plate  could  be  the  frog  Cope  described  as  brachycephala  because  Cope's  is 
western,  Yellowstone  River  is  the  type  locality,  and  Washington  State  is  the 
illustration.] 

Cope  was  indefinite  in  his  definition  of  the  ranges  of  these  three  forms.  Kauf- 
feld  gives  us  three  forms  in  the  Northeast.  We  suppose  he  thinks  of  his  Pied- 
mont R.  pipiens  much  as  we  do  of  Pseudacris  n.  feriarum,  Hyla  andersonii,  or 
Rana  virgatipes,  or  Conant's  Lampropeltis  e.  temporal  is.  The  three  frogs  above 
range  to  South  Carolina  or  Georgia  and  Conant's  snake  to  North  Carolina,  but 
Kauffeld  sends  his  jR.  pipiens  "throughout  the  Coastal  Plain  and  west  to 
Texas."  Why  does  he  not  have  it  going  up  the  Mississippi  Valley  ?  If  Cope  was 
indefinite,  Kauffeld  is  equally  so,  and  particularly  with  Rana  sphenocephala. 
He  does  not  specify  whether  it  is  an  Upper  or  Lower  Austral  or  Gulf  strip 
form.  His  arrangement  sends  three  tongues  of  meadow  frogs  into  the  south, 
brachycephala  in  the  high  Allegheny  Mountains,  ptptens  in  the  Piedmont, 
and  sphenocephala  probably  in  the  Lower  Austral  or  Gulf  strip.  This  does  not 
accord  with  our  collecting  experience  of  the  last  35  years. 

Trapido  and  Clausen  (Que.,  1038,  pp.  121-122)  tested  the  head  length  and 
tibiotarsal  lengths.  "The  remainder  of  the  characters  are  equally  unsatisfac- 
tory. .  .  .  The  material  collected  in  Quebec,  therefore,  as  well  as  specimens 
from  widely  scattered  stations  within  the  range  which  Kauffeld  has  assigned 
to  brachycephala,  fail  to  confirm  his  conclusions.  It  seems  better  for  the  present 
to  refer  the  northern  specimens  to  pipiens,  until  all  the  available  material  of 
this  widespread  and  perplexing  species  can  be  studied." 

R.  Grant  (Que.,  1944,  pp.  151-152)  finds  that  his  Quebec  meadow  frogs 
agree  with  Kauffeld's  R.  pipiens  in  head  length,  dorsal  skin  folds,  and  webbing 
of  toes.  They  agree  with  his  R.  brachycephala  in  absence  of  tympanic  spot,  in 
extension  of  dorsolateral  fold  to  supraocular  region  in  one  fourth  of  the  speci- 
mens, and  in  the  failure  of  the  tibiotarsal  joint  to  reach  the  snout  m  16  out  of 
24  specimens. 

For  a  discussion  of  Moore's  extensive  paper,  see  our  account  of  R.  p.  spheno- 
cephala. We  do  not  differ  with  him  on  pipiens  and  brachycephala  where  his  lo- 
calities are  certain,  but  his  Vermont,  Wisconsin,  and,  particularly,  southern 


FROGS:  RANIDAE  505 

material  is  not  so  certain.  His  evidence,  therefore,  against  R.  p.  sphenocephala 
is  somewhat  vitiated,  and  in  his  account  (see  R.  p.  sphenocephala)  we  see 
much  to  prove  it  a  good  form. 

Some  of  us  naturalists  have  existed  for  35  years  or  more  knowing  R.  pipiens 
and  R.  p.  sphenocephala  (R.  sphenocephala  to  Miss  Dickerson) .  Not  until  1937 
did  R.  brachycephala  ever  come  out  of  the  West  and  enter  the  eastern  horizon. 
Moore's  conclusion,  we  fear,  is  largely  against  the  three-form  conception  in- 
stead of  two  forms  for  he  writes: 

"A  study  has  been  made  of  the  taxonomic  characters  customarily  employed 
in  separating  Rana  pipiens  Schreber,  Rana  sphenocephala  (Cope)  and  Rana 
brachycephala  (Cope).  These  diagnostic  characters  were  found  invalid  when 
samples  from  many  localities  were  studied.  It  does  not  appear  possible  to  recog- 
nize three  species  or  subspecies  of  meadow  frogs  on  the  basis  of  differences  in 
body  proportions  or  pigmentation.  Therefore,  the  meadow  frogs  of  eastern 
North  America  should  be  known  as  Rana  pipiens  Schreber.  Rana  spheno- 
cephala  Cope  and  Rana  brachycephala  (Cope)  should  be  reduced  to  synonyms 
of  Rana  pipiens  Schreber."  The  case  for  R.  pipiens  and  a  supposed  eastern 
R.  brachycephala  as  one  may  be  proved  by  Moore;  but  probably  nine  of  every 
ten  naturalists  who  knew  the  meadow  frogs  of  the  East  well  have  assumed  this 
to  be  a  fact.  The  case  of  R.  p.  sphenocephala  as  thrown  into  the  scrap  basket 
is  not  a  frontal  attack  in  his  paper.  It  is  a  southpaw  affair  with  dealer's  material 
and  as  much  proves  R.  p.  sphenocephala  good  as  bad.  For  example,  in  his  argu- 
ment against  R.  p.  sphenocephala,  Moore  uses  the  tympanic  spot:  "Those  with 
sharply  defined  circular  spots  are  not  restricted  to  the  southern  states  .  .  .  , 
and  animals  from  this  region  may  lack  the  spot  entirely.  .  .  .  Thus  a  sharply 
defined  white  tympanic  spot  is  not  a  constant  feature  of  southern  populations, 
as  Kauffeld  and  others  have  suggested."  This  has  not  distressed  field  naturalists 
in  the  East  for  20  years.  Of  course  southern  Indiana,  southern  Illinois,  western 
Kentucky,  and  Oklahoma  arc  in  the  range  of  R.  p.  sphenocephala.  Moore's 
observations  help  to  prove  the  validity  of  R.  p.  sphenocephala.  Our  R.  p. 
sphenocephala  heresy  regarding  New  Jersey  has  disturbed  some  of  our  natu- 
ralists of  the  Atlantic  seaboard.  That  is  north.  But  to  push  the  case  far  toward 
the  home  of  Moore's  advisor,  who  may  rightly  believe  all  the  western  pipiens 
to  be  one,  we  quote  E.  H.  Taylor  (Kans.,  igigb,  p.  65).  In  speaking  of  the 
extreme  southwestern  corner  of  Kansas,  Morton  County  (north  not  south, 
west  not  east)  he  remarks,  "Specimens  appear  to  approach  the  characters 
of  the  southern  Rana  sphenocephala  (("ope)."  It  is  not  strange,  if  Bragg  calls 
Oklahoma  meadow  frogs  R.  p.  sphenocephala,  to  have  frogs  from  this  Kansas 
county  at  the  tip  end  of  the  Cimarron  River  drainage  (Arkansas  River  Basin) 
with  sphenocephala  tendencies.  Let  any  physical  geographer,  life-zone  en- 
thusiast (life  zones  studied  for  100  years),  or  ecologist  observe  Merriam's 
Lower  Austral  (map  i,  p.  fi),  or  Van  Dersal's  Coastal,  Piedmont,  and  Ozark 
and  Tennessee  Valley  districts  (map  2,  p.  8)  of  today,  or  Cooper's  (1859) 


506  HANDBOOK  OF  FROGS  AND  TOADS 

Carolinian,  Mississippian,  Tennessean,  and  Floridian  districts,  and  he  has  the 
natural  range  of  R.  p.  sphenoccphala,  which  follows  up  Mississippi  tributaries. 
D.  Western  puzzles. 

1.  Ranafisheri.  Plate  CVIIL 
(See  account  pp.  454-459.) 

One  cannot  help  wondering  whether  visitors  to  Las  Vegas  since  Hoover 
Dam  (Boulder  Dam)  came  have  ever  seen  fine  big-eared  males  and  other 
specimens  like  those  Mernam  and  Stejneger  had.  Since  fish  ponds  and  a  golf 
course  have  replaced  the  original  springs,  one  queries  whether  what  we  saw 
in  1925  and  what  earlier  workers  saw  are  existent  today.  Even  in  1925  the  two 
or  three  specimens  we  collected  in  the  creek  near  or  in  town  seemed  more 
like  R.  pipiens,  but  some  of  the  spring's  specimens  were  certainly  different 
from  our  New  York  R.  piptens.  Nevertheless  we  must  confess  that  it  is  of 
the  R.  piptens  complex.  Those  old  1925  specimens  were  more  distinctive  than 
the  Rana  onca  as  conceived  by  Tanner  and  Linsdale.  Bogert  and  Oliver  (Gen., 
1945,  p.  321)  hold  that  "the  relationships  of  leopard  frogs,  Rana  pipiens,  have 
been  the  subject  of  considerable  debate,  but  no  one  has  offered  convincing 
evidence  that  more  than  one  species  is  involved.  .  .  .  Nevertheless  its  failure 
to  reach  southern  California  (until  introduced)  and  the  fact  that  no  well- 
defined  races  have  evolved  (recognition  of  the  'species'  Rana  onca  and  Rana 
fisheri  to  the  contrary)  indicate  that  its  influx  into  the  region  west  of  the  con- 
tinental divide  has  been  a  recent  one." 

2.  Rana  onca.  Plate  CIX. 

Shortly  after  we  collected  R.  prettosa  with  Wilmer  W.  Tanner  at  Provo, 
Utah,  we  saw  Yarrow's  illustration  of  Rana  onca.  His  illustration  looks  like 
R.  prettosa.  We  have  never  seen  the  type  specimen.  Certainly  the  figure  looks 
little  like  the  Virgin  River  specimens  we  saw  or  like  those  from  the  same 
region  now  in  the  University  of  California  collection.  We  are  not  certain  of 
Rana  onca  as  a  good  species  or  subspecies.  Strictly,  we  suppose  we  should  dub 
it  R.  pipiens  onca. 

When  we  were  mounting  Rana  monteztimae  original  figures  from  the  US. 
and  Mex.  Boundary  Survey  and  Mission  Scientifiqtte  au  Mexique  reports  we 
noted  for  the  first  time  that  Boulenger  in  1882  placed  R.  onca  in  the  synonymy 
of  Rana  montezumne.  We  surely  cannot  blame  him  for  this  attribution  when 
even  today  we  do  not  understand  the  form  very  well  and  when  the  original 
R.  onca  figure  is  so  uncertain. 

3.  Rana  pipiens  berlandieri  (Baird).  Plate  CX. 

In  1859  Baird  described  this  form  as  follows:  "117.  Rana  berlandieri  Baird, 
Plate  56,  Figs.  7-10.  Sp.  Ch. — Size  large.  Body  stout,  robust.  Eye  distant  not 
quite  i%  times  its  diameter  from  tip  of  snout,  and  contained  2%  times  in  the 
length  of  jaw  from  rictus.  Tympanum  two-thirds  the  diameter  of  the  eye.  A 
vocal  vesicle  on  each  side  of  the  head.  A  glandular  fold  on  each  side  the  jaw, 
and  another  one  broad  and  depressed  on  each  side  of  the  body.  Between  these 


Plate  CVHL  Rana  fisheri. 


Ncv- 


508 


HANDBOOK  OF  FROGS  AND  TOADS 


is  one  pair  of  ridges  along  the  coccyx;  several  pairs  more  interrupted  anterior 
to  it.  Skin  corrugated  and  irregular,  quite  pustular  in  some  specimens.  Feet 
webbed  from  the  bulb  of  the  toes;  excavated  on  the  inner  edges;  last  joint  of 
longest  free.  Femur  about  half  the  length  of  body,  shorter  than  the  tibia.  Color 
above  greenish  olive,  with  distant  subcircular  blotches  of  darker,  scarcely 
areolated  in  the  preserved  specimens.  Beneath  yellowish  white,  with  brown 
mottlings  on  the  throat.  An  indistinct  whitish  line  on  the  side  of  the  head; 
especially  jn  the  young;  the  lateral  ridges  bronzed.  Southern  Texas  generally" 
(US.  and  Mcx.  Bound.  Survey,  vol.  II,  pt.  2,  pp.  27-28). 


Plate  CIX.  Rana  onca.  i.  From  Geog.  Survey  E.  of  looth  Meridian,  Plate 
XXV.  2,3.  From  2  miles  southeast  of  Overton,  Nev.  (X%). 

For  some  time  this  form  was  quite  universally  used  and  its  range  inter- 
preted as  widespread,  not  southern  Texas  alone.  Yarrow  (Ariz.,  1875,  p.  527) 
had  it  from  Provo,  Ephraim  City,  and  Beaver,  Utah,  to  Denver,  Colo.,  south 
to  six  localities  in  northern  New  Mexico  (much  of  New  Mexico  and  much 
of  Colorado  tributary  to  the  Rio  Grande  River),  In  1878  (Mont.,  p.  289),  Coues 
and  Yarrow  (Dakotas  and  Montana)  extended  it  to  the  Pembina  Mountains 
with  this  disclaimer:  "The  common  Western  form  is  Rana  halecina  berlandieri 
which  is  only  distinguished  from  R.  halecina  by  its  larger  size  and  generally 
coarser  and  more  pustulated  skin.  The  specimens  represent  'berlandieri'  but 
this  we  are  disinclined  to  adopt  without  further  investigation  of  its  alleged 
distinctness." 

In  1880  (Tex.,  i88oa,  p.  24)  Cope  had  "Rana  halecina  berlandieri  Bd.  Com- 
mon at  Dallas  and  on  the  first  plateau;  also  in  the  low  country  near  Wash- 
ington on  the  Brazos" — i.e.,  in  northern  Texas  outside  the  Rio  Grande  Valley. 
In  fact,  in  1879  Cope  (Mont.,  p.  436)  called  Rana  halecina  berlandieri  the 
abundant  species  on  the  plains,  i.e.,  he  had  it  1879-1880  from  the  Rio  Grande 


FROGS:  RANIDAE  509 

to  Montana.  But  in  1889  he  described  Rana  virescens  brachycephda  for 
meadow  frogs  from  Provo,  Utah,  to  Framingham,  Mass.,  from  Montana  to 
Chihuahua,  and  the  second  frog  on  his  list  is  from  Brownsville,  near  or  at  the 
actual  home  of  R.  bcrlandicri. 

If  the  name  is  employed,  we  would  restrict  it  to  southern  Texas  or  the  Rio 
Grande  Valley  until  it  is  better  described  and  understood.  This  position  ap- 
proaches that  of  Schmidt  (1944,  p.  81),  who  holds,  "The  name  berlandten 
appears  available  for  the  large  form  of  Rana  pipiens  found  at  Brownsville, 
and  it  is  reasonable  to  suppose  that  all  the  leopard  frogs  of  the  Rio  Grande 
drainage  are  derived  from  this  center.  ...  In  the  present  state  of  the  pipiens 
problem,  it  is  preferred  to  employ  the  earliest  name  available  that  is  reasonably 
well  associated  geographically  with  our  area." 

Like  K.  P.  Schmidt  and  T.  F.  Smith  (Tex.,  1944),  M.  B.  Mittleman  and 
H.  T.  Gier  (Gen.,  1942,  pp.  7-15)  feel  the  emphasis  should  be  on  the  western 
forms. 

"Throughout  the  several  discussions  briefly  mentioned  above,  there  is  only 
scant  mention  of  Baird's  Rana  berlamUcn  (1859:  27)  described  on  the  basis 
of  eleven  specimens  from  'Southern  Texas  generally.'  We  find  in  extensive 
series  of  leopard  frogs  from  Texas,  Kansas,  Oklahoma  and  possibly  certain 
adjacent  States,  characteristics  which  are  not  in  accord  with  equivalent  series 
of  specimens  from  the  Atlantic  coastal  plain,  the  Appalachian  highlands  or 
more  northerly  sections  of  the  mid-western  United  States  or  from  the  Rocky 
Mountains  and  Pacific  States.  There  is  wide  variation  in  the  Texas-Kansas- 
Oklahoma  series,  as  is  natural  with  frogs  of  this  group;  none  the  less,  it  is 
comparatively  easy  to  select  specimens  from  these  three  states  no  matter 
whether  they  are  mixed  with  examples  from  other  areas  or  encountered  alone. 
Comparison  of  specimens  from  the  three  States  mentioned,  with  the  cotypes 
of  Baird's  berlandien,  shows  immediately  that  they  arc  syntypical"  (Mittleman 
and  Gier,  Gen.,  1942,  p.  9). 

In  view  of  the  fact  that  several  Texas  forms  are  now  known  to  extend  into 
Oklahoma  (Pseudacns  strccf{en  and  Scaphtopus  h.  hnrtent)  or  even  into 
Kansas  (Microhyla  olivacea  and  P.  n.  clartyi)*  there  is  considerable  evidence 
to  suggest  this  grouping.  In  1943  Stejncger  and  Barbour  accepted  this  inter- 
pretation, but  in  1943  A.  N.  Bragg  and  C.  C.  Smith  (Okla.,  194$,  p.  107)  called 
all  their  Oklahoma  meadow  frogs  /?.  sphcnocephala  Cope.  They  stated:  "We 
are  not  ready  to  express  an  opinion  as  to  the  correct  name  of  this  frog.  Mittle- 
man and  Gier  (1942)  call  it  R.  ptpiens  berlandien  (Baird)."  We  will  follow 
A.  N.  Bragg,  who  has  seen  specimens  from  all  except  nine  counties  of  Okla- 
homa. Who  knows  but  that  Texas  has  three  meadow-frog  populations — 
northeastern,  coastal,  and  forest  R.  p.  sphcnocephala;  Rio  Grande  Valley 
R.  p.  bcrlandtcri;  and  a  form  showing  northern  influence  found  on  a  prairie 
central  Texas  tongue  extending  southward  through  the  Panhandle.  (See  our 
R.  p.  berlandien  plate.)  Not  until  we  are  farther  along  in  careful  local  state 


HANDBOOK  OP  FROGS  AND  TOADS 

studies  like  Bragg's  will  we  venture  a  positive  opinion.  Our  inclination  is  to 
make  the  provisional  R.  p.  berlandieri  southern  Texas  to  San  Antonio,  i.e., 
unite  the  Nueces,  San  Antonio,  and  possibly  the  Guadalupe  rivers  with  the 
Rio  Grande  drainage— roughly  San  Antonio  to  the  southeastern  corner  of 
New  Mexico,  most  of  that  state  to  southern  Colorado.  One  look  at  the  original 
figure  and  our  Beeville  and  our  San  Antonio  pictures  show  considerable 
diversity,  possibly  three  influences  (Plate  CX). 


Plate  CX.  i.  Runa  pipiens.  From  Helotes,  Tex.  2.  Rana  ptpicns.  From  Beeville, 
Tex.  3,4,5.  R.  berlandieri.  From  U.S.  Boundary  Survey,  Plate  36.  (X%). 

* 

Mittleman  and  Gier  consider  Floridian  and  extreme  southeastern  states  as 
R.  sphenocephala  (not  the  Mississippi  Valley  to  the  Wabash  Valley  and  all  of 
Oklahoma).  For  their  comparisons  with  their  Texas-Oklahoma-Kansas  R. 
berlandieri  they  use  all  the  rest  except  "the  debatable  coastal  strip  from  South- 
ern Quebec  to  southern  New  Jersey"  as  R.  brachycephala.  We  do  not  know 
any  such  coastal  strip.  Bragg's  R.  p.  sphenocephala  cuts  in  two  their  R.  ber- 
landieri range.  If  one  were  to  recognize  a  northeastern  race  of  pipiens,  a  west- 
ern and  northwestern  race  of  brachycephala,  a  southern  and  Mississippi  Valley 


FROGS:  RANIDAE 

race  of  sphenocephala,  and  a  Rio  Grande  Valley  race  (to  southern  Colorado), 
he  would  have  in  these  populations  elements  of  all  four  to  compare  with  their 
Kansas-Oklahoma-Texas  race. 

Moore  (1944,  p.  349)  wrote:  "Recently  Mittleman  and  Gier  (1942)  have 
suggested  that  the  meadow  frogs  of  'Texas,  Kansas  and  Oklahoma  and  pos- 
sibly certain  adjacent  States'  are  distinct  from  other  populations  and  propose 
to  call  them  Rana  pipiens  berlandieri.  Their  evidence  was  given  careful  con- 
sideration, but  no  support  for  their  contention  could  be  gathered  from  the 
living  and  preserved  material  used  in  this  report." 
4.  Rana  pipiens  brachycephala.  Plate  CXI. 


Plate  CXI.  R.  pipiens  (brachycephala) .  Left.  From  Larry  Flower,  Elko,  Ncv. 
Right.  From  Deeth,  Nev. 

In  1889  Cope  described  R.  pipiens  brachycephala.  In  his  69  lots  of  188  speci- 
mens his  second  number,  3293,  consisted  of  7  specimens  from  Brownsville, 
Capt.  S.  Van  Vliet,  United  States  collector.  This  may  mean  he  considered 
R.  berlandieri  Baird  of  1859  as  a  synonym.  He  put  it  in  the  synonymy  as  Rana 
halecina  berlandieri  Cope  Check  List  Batr.  Rept.  N.  Amer.,  p.  32;  nee.  Rana 
berlandieri  Baird.  The  only  specimens  of  Rana  virescens  virescens  Cope  had 
from  Texas  were  a  Dallas  specimen  and  one  lot  of  TO  from  Matnmoros,  Mexico. 
These  may  have  been  in  the  collection  bought  from  Berlandier  at  Matamoros. 
Did  they  come  from  Matamoros?  Cope's  type  no.  3363  came  from  the  Yellow- 
stone River  and  he  figured  a  Fort  Walla  Walla,  Wash.,  specimen  no.  10922. 
Of  his  69  lots,  63  came  from  Illinois  or  Wisconsin  westward,  or  181  of  the  188 
specimens.  Of  these  181  specimens  only  3  came  from  Iowa,  2  from  Illinois, 
2  from  Wisconsin,  and  i  from  Minnesota.  This  means  only  4  east  of  the 


512  HANDBOOK  OF  FROGS  AND  TOADS 

Mississippi.  These  4  frogs  with  6  lots  (7  specimens)  from  east  of  Mississippi 
make  11  individuals  of  the  188.  The  6  lots  are:  2  Maine,  i  Massachusetts,  i 
Connecticut,  i  Quebec,  i  South  Carolina,  i  Michigan. 

Said  Cope  (Gen.,  1889,  p.  405) :  "This  is  the  common  and  only  species  of 
Rana  found  between  the  eastern  part  of  the  Great  Plains  and  the  Sierra  Nevada 
Mountains.  It  is  common  wherever  there  is  sufficient  water  to  supply  its  neces- 
sities. In  some  of  the  Western  towns  it  is  eaten  in  the  restaurants,  and  I  have 
not  infrequently  found  it  excellent  food  when  the  larder  of  my  expeditions 
in  search  of  fossils  has  run  low." 

Before  we  give  you  his  original  description  we  cannot  resist  the  observation 
that  we  believe  R.  p.  brachycephala  is  western  if  it  is  anything  at  all. 

"The  muzzle  is  less  elongate,  and  the  extended  hind  leg  brings  the  heel  to 
its  apex,  but  not  beyond.  The  tympanic  disk  is  two-thirds  the  diameter  of  the 
eye.  The  head  is  shorter,  entering  the  length  of  the  head  and  body  three  and 
a  half  times.  The  dorsal  dermal  plicae  are  thicker  and  there  are  but  two  be- 
tween the  dorsolaterals;  usually,  however  there  are  four,  as  in  the  other  sub- 
species. First  finger  longer  than  second.  Web  leaving  two  free  phalanges  of 
the  fourth  digit,  but  so  repand  as  to  give  the  antepenultimate  phalange  only 
a  wide  border.  The  inner  cuneiform  tubercle  is  rather  small,  but  has  a  rather 
prominent  compressed  edge.  External  tubercle,  none.  A  thick  tarsal  fold.  There 
are  no  large  warts  on  the  skin,  but  there  are  occasionally  minute  warts  and 
folds  on  the  superior  face  of  the  tibia. 

"In  life  the  color  of  the  superior  surfaces  is  green.  The  dorsolateral  ridges  are 
light  yellow,  and  so  is  a  stripe  from  the  end  of  the  muzzle,  which  passes  above 
the  lip  and  below  the  eye  and  tympanum  to  above  the  middle  of  the  humerus. 
There  are  two  rows  of  large  rounded  dorsal  spots  between  the  dorsolateral 
ridges,  which  are  edged  with  greenish-yellow.  There  are  two  similar  rows 
on  each  side,  of  which  the  inferior  is  the  smaller,  which  are  not  regularly  ar- 
ranged. There  is  a  spot  on  each  eyelid  and  one  on  the  end  of  the  nose  above. 
There  is  a  light  band,  frequently  broken  into  spots  near  the  edge  of  the  upper 
lip.  There  is  a  brown  spot  on  the  elbow  and  one  on  the  front  of  the  cubitus. 
The  band  seen  on  the  front  of  the  humerus  in  R.  v.  virescens  is  here  an  illy 
defined  spot.  On  the  superior  face  of  the  femur  there  are  three  brown  spots, 
but  there  is  no  longitudinal  brown  band  in  front  of  these  spots,  as  is  usual  in 
the  two  other  subspecies  of  the  R.  viresccns.  There  arc  three  complete*  wide 
brown  cross-bands  on  the  femur,  and  sometimes  four.  Frequently  there  are 
one  or  two  spots  on  one  or  both  faces  of  the  tibia  besides  the  three  bands.  The 
posterior  face  of  the  femur  is  greenish  llow,  coarsely  marbled  with  brown. 
All  the  spots  and  bands  are  narrowly  y  iw-edged.  Inferior  surfaces  are  light 
yellow,  unspotted"  (E.  D.  Cope,  Gen.,  9,  pp.  403-404) . 

Note  the  last  sentence. 

Two  years  after  this  description  appe  d,  Dr.  Stejneger  (Calif.,  1893,  p.  228) 
used  Rana  pipiens  brachycephala  for  I  aranagat  Valley,  Nev.,  material.  The 


FROGS:  RANIDAE  513 

same  year  (Ind.,  1892,  p.  474)  O.  P.  Hay  characterized  this  form,  ending, 
"Maine  to  Oregon  and  Mexico,  but  mostly  western  U.S.  .  .  .  The  variety 
sphenocephala  is  chiefly  southern  in  its  range,  pipiens  (vtrescens  of  Cope) 
eastern  and  northern,  and  brachycephala  western.  Neither  is  confined,  how- 
ever, to  these  limits.  With  us  the  common  form  is  pipiens,  and  it  is  every- 
where abundant.  Sphenocephala  is  in  the  National  Museum  from  Wheatland, 
Ind.,  and  to  this  I  refer  one  specimen  from  Lake  Maxinkuckee,  which  I  find  in 
the  collection  of  the  State  Normal  School.  One  specimen  in  the  same  collec- 
tion and  taken  at  Camden,  Carroll  County,  had  some  of  the  characteristics  of 
brachycephala.'' 

Kellogg  has  written  about  R.  h.  berlundien  Cope  as  follows:  "Until  the 
publication  of  Cope's  'Batrachia  of  North  America,'  the  subspecific  name 
brachycephala  was  strictly  a  nomen  midiim,  inasmuch  as  the  characters  that 
distinguished  this  subspecies  from  supposed  typical  hnlectna  (vtrescens  Cope, 
1889)  were  not  indicated.  Cope's  diagnosis  of  the  subspecies  brachycephala 
published  in  1889  is  seemingly  based  upon  one  specimen,  and  this  inference 
is  drawn  from  his  introductory  remark  that  4I  select  as  typical  a  specimen  from 
the  Yellowstone  River  (no.  3365).'  There  are,  however,  15  specimens  cata- 
logued under  that  number,  although  Cope  mentions  only  5  in  his  list  of  speci- 
mens examined,  all  of  them  were  collected  by  Dr.  F.  U.  Hayden  in  1857  along 
the  Yellowstone  River  in  Montana.  This  lot  consists  of  2  adults,  i  young  and 
12  tadpoles.  Inasmuch  as  it  is  now  impossible  to  recognize  the  actual  individual 
upon  which  Cope  based  his  description,  all  of  these  specimens  have  been  desig- 
nated as  the  cotypes  of  the  subspecies  brachycephala.  The  following  notes  apply 
to  the  largest  cotype:  Head-and-body  length,  86;  transverse  diameter  of  tympa- 
num, 6;  transverse  diameter  of  eye,  8;  anterior  edge  of  eye  to  nostril,  5.4  mm.; 
the  hind  limb  being  carried  forward  along  the  body,  the  tibio-tarsal  joint 
reaches  to  the  anterior  margin  of  eye"  (Kellogg,  Gen.,  1932,  pp.  208-209). 

If,  sometime  in  the  future,  students  prove  that  the  western  and  northwestern 
form  is  different  from  the  northeastern,  then  we  would  recommend  R.  p. 
brachycephala.  We  must  confess  that  in  1925,  1954,  and  T942>  we  did  not  note 
any  surprising  differences  from  R.  pipiens  proper  in  the  northern  Nevada, 
Utah,  or  Wyoming  meadow  frogs.  In  1942,  for  example,  we  made  these  notes: 

May  22,  1942,  Deeth,  Nev.  Short  distance  beyond  is  pool  with  bushes.  Bert 
caught  meadow  frogs  here,  two  females  ripe,  much  bigger  than  Virgin  River 
frogs  and  dorsal  spots  larger  and  very  light-margined. 

May  23,  Elko,  Nev.  Went  to  Maggie  and  Susie  creeks  at  Carlin.  On  way 
back  saw  near  Humboldt  River  in  overflow  six  Rana  pipiens  with  numerous 
dorsal  spots  and  these  light-encircled.  Caught  three,  one  male  and  two  half- 
grown. 

May  30.  With  Wilmer  Webster  Tanner,  went  i  mile  west  of  Provo  to  a 
spring.  Beside  the  stream  took  a  R.  pipiens. 

June  5,  north  of  Kanab.  A  fine  pond,  fish  in  it— very  quagmirelike.  Saw 


514  HANDBOOK  OF  FROGS  AND  TOADS 

four  meadow  frogs.  Caught  three.  Meadow  frogs  out  of  water  were  in  a 
meadowhke  area. 

These  notes  reveal  no  feeling  of  differences  on  our  part.  We  are  reminded 
that  in  1915  Ruthven  and  Gaige  in  northern  Nevada  held,  "The  specimens 
show  little  variation  from  Dickerson's  (The  Frog  Boof(,  171)  description." 

Recently,  almost  everyone  has  called  this  form  Rana  ptptens.  We  are  glad 
to  see  that  when  A.  and  R.  D.  Svihla  (Wash.,  1933,  p.  125)  do  use  a  subspecies 
term  they  employ  R.  p.  brachycephala. 
5.  Arizona  problems. 

After  studies  in  the  Navaho  country  or  Painted  Desert  region  T.  H.  Eaton, 
Jr.  (Ariz.,  1935,  p.  150)  wrote,  "The  great  variability  of  ptpiens  in  the  south- 
west, appears  ample  reason  for  doubting  the  validity  of  onca,  since  numerous 
intermediates  occur."  This  is  true  not  only  of  southern  Nevada  and  southern 
Utah  but  also  of  all  Arizona.  Witness  the  following  notes  made  at  various 
places  in  Arizona  where  it  seemed  as  if  each  spring  or  moist  area  had  a  dif- 
ferent form, 
a  (i).  Anvaca  and  Pena  Blanca  Springs. 

June  17, 1934,  went  with  Prof,  and  Mrs.  L.  P.  Wehrlc  and  family  to  Arivaca. 
Here  is  a  fine  stream  with  water  now  in  pools — all  along  it  are  willows  and 
huge  cottonwoods.  In  one  pool  found  tadpoles  of  R.  pipicns  and  some  trans- 
formed ones.  Secured  a  fine  male  and  female  of  Rana  pipiens.  They  are  so 
different  from  ours  in  Ithaca,  New  York! 

June  18,  Pena  Blanca  Spring.  Came  here  Monday.  Arrived  at  dusk.  In  the 
overflow  of  the  spring  one  meadow  frog  and  one  Hyla  arenicolor—one 
meadow  frog  in  cow  trough.  They  are  much  spotted  and  yellow  under  legs 
like  the  Arivaca  ones. 

June  21,  Pena  Blanca.  Went  with  Mr.  G.  W.  Harvey.  In  the  shallower  pools 
found  some  R.  pipiens  adults,  young,  and  tadpoles  as  well.  Most  of  the  big 
rocky  pools  had  only  R.  tarahumarae  adults  and  tadpoles,  while  the  more 
shallow  ones  had  R.  pipiens. 

June  25,  1942.  Today  we  went  to  Arivaca.  Went  the  length  of  this  creek 
and  along  irrigation  ditch.  Saw  plenty  of  meadow  frogs, 
a  (2).  Fauna  of  Pajaritos  Mts. 

".  .  .  Of  batrachians,  a  toad  (Bufo)  and  a  frog  (Rana  virescens  brachy- 
cephala Cope)  were  found  at  Warsaw  Mills;  and  at  Buenos  Ayres,  at  the  be- 
ginning of  the  summer  rams  .  .  ."  (Mearns,  Ariz.,  1907,  p.  113). 

(This  locality  like  Arivaca  and  Pena  Blanca  Springs  is  close  to  the  Mexican 
boundary  and  just  northwest  of  Nogales.) 
b.  Sabino  Canyon. 

July  2,  1934.  We  went  about  5  to  picnic  spots  where  there  were  two  pools. 
The  upper  pool  was  filled  with  Rana  pipiens.  They  would  often  climb  up  on 
slanting  rocks,  some  8-10  feet,  possibly  for  insects.  They  are  very  orange  on 
underside  of  legs. 


FROGS:  RANIDAE 


515 


When  we  observed  the  decided  orange  color  on  the  ventral  surface  of  the 
hind  legs,  we  did  not  know  that  King  had  anticipated  us  by  2  years.  He  had 
written : 

"Leopard  frogs  were  common  along  the  stream  in  Sabino  Canyon.  One 
large  individual  was  also  found  under  a  bunch  of  hay  on  University  Farm,  at 
least  two  hundred  yards  from  any  permanent  water.  The  young  were  trans- 
forming the  latter  part  of  July.  All  individuals  examined  showed  very  bright 
orange  coloration  on  backs  of  thighs  and  on  flanks.  The  dorsal  spotting  is  ir- 
regular; the  tympanum  may  or  may  not  have  a  light  spot;  the  spot  on  the 
snout  is  also  variable;  the  legs  are  barred.  The  general  coloration  is  grayish 
with  many  showing  green  or  brown  as  a  dominant  color"  (F.  W.  King,  Ariz., 
1932,  p.  176). 
c.  Carr  Canyon,  Huachuca  Mts.  Plates  CXII;  CXIIL 


Plate  CXII.  Rana  piptens  (X1/^)-  Females,  from  Carr  Canyon,  Ariz. 

July  i,  1942.  On  a  bank  of  the  reservoir  I  espied  four  fat  spotted  frogs.  I  made 
a  pass  at  one  and  it  slipped  out  of  my  hands.  The  others  were  frightened.  It 
was  a  precarious  spot.  Then  Anna  suggested  I  try  the  fish  pole  noose  on  them. 
One  was  out  of  water  on  the  bank.  In  spite  of  the  fact  that  the  noose  could  be 
brought  only  to  the  shoulders  we  were  both  amazed  at  the  results.  The  first 
one  leaped  through  the  noose,  the  next  one  on  the  bank  was  captured,  and  how 
Anna  did  laugh  at  the  success  of  the  method.  The  third  stuck  its  head  and  the 
forepart  of  its  body  out  of  the  water  and  I  draped  the  noose  around  its  neck 
with  no  concern  on  its  part.  This  one  I  also  caught.  These  are  large  frogs. 

Color:  The  ground  color  of  back  is  drab-gray  or  light  drab.  The  spots  are 
black  or  fuscous  with  centers  of  drab  and  slight  thread  of  pale  vinaceous-fawn 
outlining  them.  The  sides  are  avellaneous,  the  groin  marbled.  The  dorsolateral 
folds  are  drab-gray  or  light  drab.  There  are  five  or  six  broken  folds  on  back 


5i6  HANDBOOK  OF  FROGS  AND  TOADS 

between  the  dorsolateral  ones  and  also  one  or  two  on  leg.  The  bands  on  legs 
and  tarsus  are  drab.  The  lower  jaw  is  mottled.  There  is  no  pronounced  light 
center  in  tympanum.  Under  parts  are  in  part  white  or  ivory  yellow,  with  the 
underside  of  legs,  femur,  tibia,  and  top  of  tarsus  and  foot  antimony  yellow, 
with  a  slight  wash  of  same  in  axil  of  arm,  on  brachium,  lower  belly,  and  groin. 

The  measurements  in  millimeters  of  two  males  follow:  length  (snout  to 
vent)  88,  89;  head  (tympanum)  28,  27;  head  (angle  of  mouth)  26,  25;  width 
of  head  29,  29;  snout  13,  13;  eye  14,  14;  mterorbital  space  4,  4.5;  upper  eyelid 
5,  5.5;  tympanum  7.5,  6;  mtertyrnpanic  space  20.5,  19.5;  mternasal  space  7,  n; 
fore  limb  32,  45;  first  finger  13,  13;  second  finger  12,  12.5;  third  finger  19.5, 
18.5;  fourth  finger  14,  14.5;  hind  limb  124,  134;  tibia  48,  47;  foot  with  tarsus 
60,  60;  foot  without  tarsus  41,  41;  first  toe  n,  15;  second  toe  20,  22;  third  toe 
28,  }2;  fourth  toe  39.5,  42;  fifth  toe  27,  29.5;  hind  limb  from  vent  134,  155. 

When  we  came  to  the  deep  pool  6-ro  feet  below  us  in  Carr  Canyon,  we  be- 
held what  did  not  look  like  R.  pipiens.  Were  we  seeing  R.  forren  or  R.  omil- 
temana  of  the  Biologia  Centrah-Americana  figures  or  the  predecessors  of 
R.  areolata  of  the  United  States  or  Rana  montezumae  or  R.  p.  berlandien* 
We  could  get  them  alive  only  by  noosing. 

By  Boulenger's  1920  key  the  frog  traces  to  Rana  montezumae.  With  Kellogg 
(1932)  we  get  the  same  result.  If  by  these  two  conservative  authors  we  arrive 
at  R.  montezumae,  it  is  apparent  that  the  Huachuca  Mountain  frogs  need 
to  be  carefully  examined  with  a  larger  series.  We  have  not  compared  it  with 
good  new  R.  montezumae  material  from  mid-Mexico.  We  keep  this  form  in 
the  R.  pipiens  complex  until  more  data  appear.  By  measurements  and  white 
stippling,  however,  it  appears  to  be  R.  montezumae. 

In  their  description  of  the  type  of  7?.  areolata,  S.  F.  Baird  and  C.  Girard 
(Gen.,  1852,  p.  173)  remarked  that  they  had  "a  very  small  one  on  the  Rio  San 
Pedro  of  the  Gila."  We  wonder  what  their  frog  was. 
d.  Lakeside,  Ariz.,  Plate  CXIV;  CXV. 

July  9,  1942.  Show-Low,  Ariz.  Stopped  at  1:30  at  Carrizo  Creek,  Gila  Co., 
Ariz.  Saw  a  meadow  frog  near  edge  of  pool.  Caught  it.  Two  more  jumped  in. 
Near  another  pool  three  more  leaped  in.  In  Chara  found  fish,  damsel  fly,  and 
meadow  frog  tadpoles.  Seemingly  straight  R.  pipiens. 

July  9.  Lakeside,  Ariz.,  on  the  edge  of  Mogollon  Mesa.  One  mile  north  of 
Lakeside,  Ariz.,  saw  an  interesting  pool  beside  the  road  with  no  ei\d  of 
meadow  frogs.  Put  up  at  Lake-O- Woods  Lodge.  Went  down  to  a  swampy 
area  below  the  swimming  pool  dam,  and  found  countless  puzzling  meadow 
frogs.  Went  to  a  drainage  or  irrigation  ditch  which  goes  off  from  the  swim- 
ming pool  pond.  Here  along  the  steep  walls  saw  some  frogs  which  at  first 
looked  like  bullfrogs  or  Rana  tarahumarae,  and  some  were  surely  Rana 
pipiens  in  appearance.  Tonight  early  in  evening  heard  no  frogs. 

July  10.  In  the  irrigation  ditch  cut  through  rock  we  noosed  several  of  the 
large  frogs.  One  was  brown  with  no  spots;  another,  a  male,  was  green  with 


FROGS:  RANIDAE 


517 


very  little  indication  of  spots;  and  another  was  quite  R.  pipiens-Yike.  These 
frogs  are  a  queer  lot.  Are  they  all  one?  Some  were  quite  wary  and  would 
sink  into  vegetation  as  the  end  of  pole  and  noose  approached.  We  went  down 
to  lower  pond.  Caught  frogs  from  transformation  to  full  grown.  Many  were 
caught  with  the  noose.  How  they  would  leap  at  the  line!  As  we  came  along 
the  edge  of  the  lake  and  waded  out  to  catch  frogs  out  of  the  floating  mat 
with  our  fish  pole  noose  we  discovered  four  masses  of  eggs  near  the  hank. 


Plate  CXllL  Left.  Rana  Iccontci.  From  Mission  Sc.  du.  Mexique,  Plate  4, 
Fig.  i.  Center.  Rana  jorrert.  From  Bwl.  Centr.  Amer.,  Plate  60,  Fig.  A.  Right. 
Rana  omiltemana.  From  BioL  Centr.  Amer.,  Plate  61,  Fig.  A. 


Plate  CXIV.  T.  Rana  pustttlosa.  BioL  Centr.  Amcr.,  Plate  61,  Fig.  B.  2,3.  Rana 
montezumae.  From  U.S.  and  Mex.  Boundary  Survey,  Plate  36,  Figs.  1,5. 

They  are  plinths  attached  to  milfoillike  plants.  Presently  near  bank  found  a 
fresh  mass.  Were  they  laid  when  rain  threatened  last  night?  The  other  four 
masses  were  approaching  hatching.  A  few  days  ago,  about  Thursday  of  last 
week,  a  small  rain  came.  The  uniform  frog  now  has  spots  on  its  back  like 
1?.  pipiens.  The  green  bullfroglike  one  has  a  few  spots. 

July  ii.  Today  Anna  and  I  went  along  the  drainage  ditch  west  of  the  lower 
pond.  Here  saw  many  snakes.  Plenty  of  frogs  in  the  ditch.  Saw  no  tadpoles. 

July  13.  At  S,500-ft.  elevation,  east  of  McNary,  20  miles  or  more,  on  Eager 
side  of  crest  found  a  fine  spring  with  many  Rana  pipiens.  They  look  different 
from  Lakeside  frogs,  so  also  the  tadpoles. 

Color:  Lakeside,  Ariz.  Large  female.  The  background  between  costal  folds 


X 


I 


FROGS:  RANIDAE  519 

is  grass  green,  sides  sayal-brown  or  mikado  brown  with  rear  of  eyelid  the 
same,  and  upper  jaw  grass  green.  Between  the  dorsolateral  folds  are  bare 
suggestions  of  other  folds.  The  top  of  fore  and  hind  limbs  is  bister.  Down  the 
back  are  3  rows  of  large  but  obscure  cinnamon-brown  spots.  Below  the  lateral 
folds  are  2  or  3  rows  of  very  small  spots  and  here  also  are  many  tubercles.  The 
rear  of  back  has  same,  also  rear  of  fore  limbs,  and  top  of  hind  limbs.  The  tibia 
has  suggestion  of  3  or  4  tubercular  ridges.  One  tibial  ridge  of  tubercles  runs 
along  foot  almost  to  tip  of  fifth  toe,  and  an  inner  tarsal  ridge  goes  to  metatarsal 
tubercle.  The  groin  is  mottled  with  cinnamon-brown  or  black  and  antimony 
yellow.  There  are  4  half-bars  on  femur  and  2  full  and  3  broken  bars  on  tibia, 
2  prominent  bars  on  tarsus,  and  6  on  foot  counting  down  the  fourth  toe.  The 
rear  of  femur  is  black,  spotted  with  rough  tubercles,  which  are  antimony 
yellow  or  yellow  ocher.  The  underside  of  hind  legs  and  top  of  foot  are  mustard 
yellow.  The  under  parts  of  body  bear  wash  of  same  or  of  naples  yellow.  The 
throat  and  underside  of  fore  limbs  are  pale  ochraccous-buff.  The  iris  is  bister 
with  light  vinaceous-cinnamon  streaks  and  clear  dull  green-yellow  on  outer 
edge  of  eye. 

Male.  The  back  is  olive-green,  becoming  on  the  sides  pyrite  yellow.  Spots 
are  very  obscure,  barely  showing,  black.  The  upper  jaw  from  snout  to  tympa- 
num is  lettuce  green.  There  are  prominent  tubercles  below  the  lateral  folds 
and  some  between  them.  The  whole  rear  of  legs  and  back  of  fore  limbs  are 
tubercular,  the  tibia  with  3  or  4  tubercular  ridges.  The  belly  is  white,  with 
underside  of  throat  cream  color  clouded  with  lettuce  green,  thus  reminding 
one  of  the  bullfrog.  Under  legs  and  groin  are  mustard  yellow  as  in  female. 
This  frog  has  no  lateral  vocal  sacs. 

A  young  frog  (smaller  than  typically  spotted  pipiens)  has  a  bright-green 
mask  and  green  back  and  is  colored  just  like  the  male  described  above. 

Small  frog.  Background  of  back  fawn,  becoming  below  dorsolateral  folds 
vmaceous-fawn.  Very  few  tubercles  arc  apparent.  There  are  four  prominent 
ridges  between  the  dorsolateral  folds.  These  folds  and  the  one  on  upper  jaw 
are  deep  olive-buff  or  olive-buff.  The  dorsal  spots  of  body  and  legs  are  black 
outlined  with  tilleul  buff.  Upper  side  of  fore  limbs  is  vmaceous-fawn.  Under 
parts  are  entirely  white  except  for  a  little  yellow  cm  groin  and  foot. 

Comments: 

(1)  Except  for  their  much  greater  size  these  frogs  at  first  reminded  us 
of  our  1925  impressions  ot  R.  fishen  at  the  original  springs  of  Las  Vegas,  Nev, 

(2)  Some  specimens  at  first  and  at  a  distance  suggested  Rana  clanntans. 

(3)  Are  they  a  part  of  the  unstable  population  (northern  Arizona  and 
southern  Utah)  from  which  Tanner's  and  Linsdale's  R.  onca  came? 

(4)  Were  the  frogs  of  the  lower  pond  (Rana  fipieni)  the  same  as  the 
forms  from  the  rock-walled,  canyonlike  irrigation  ditch ?  The  tadpole  tails 
with  heavier  spots  (although  of  the  R.  ptpiens  class)  suggest  areolata  tad- 
pole coloration. 


520  HANDBOOK  OF  FROGS  AND  TOADS 

(5)  Were  these  colorations  somewhat  akin  to  our  northeastern  dark 
jR.  pipiens  just  out  of  hibernation,  and  were  these  frogs  only  a  short  time  out 
of  hibernation? 

Western  Spotted  Frog,  Western  Frog,  Pacific  Frog,  Spotted  Frog 

Rana  prctiosa  pretiosa  (Baird  and  Girard).  Plate  CXVI;  Maps  30,  35. 

Range:  Northern  California  to  southeastern  Alaska,  and  east  to  Waskesm 
Lake,  Saskatchewan,  Montana,  and  western  Wyoming.  Through  central  Utah 
to  northern  Arizona  (Tanner  and  Hunt). 

"In  southern  British  Columbia  Rana  pretiosa  and  Rana  aurora  occupy 
geographically  complementary  ranges.  Thus  prctiosa  is  recorded  from  many 
localities  east  of  the  coast  range  and  reaches  the  coast  north  of  Prince  Rupert, 
while  aurora  is  known  only  from  the  coast  of  extreme  southwestern  British 
Columbia  and  the  adjacent  islands.  Logier  .  .  .  cites  a  record  of  pretiosa 
on  Sumas  Prairie  in  the  center  of  the  area  occupied  by  aurora.  It  is  significant 
to  record  the  capture  of  two  recently  transformed  specimens  of  ptetiosa  on 
Nicomen  Island,  in  the  Fraser  River  some  50  miles  east  of  Vancouver,  on 
October  20,  1941.  The  occurrence  of  these  two  species  on  the  same  general 
territory  provides  an  opportunity  to  investigate  their  ecological  relations" 
(G.  C.  Carl,  B.C.,  1945,  p.  53). 

Habitat:  Aquatic.  In  pools  and  marshes  along  permanent  streams,  in  lakes 
or  springs  of  mountainous  sections. 

"Common  along  streams,  but  nowhere  plentiful.  A  stupid  frog,  easily 
caught;  neither  a  strong  jumper  nor  a  fast  swimmer.  The  salmon  color  of 
the  underside  is  absent  from  the  newly  transformed  adult;  it  increases  m  ex- 
tent and  brilliancy  with  increase  in  si/e,  occasionally  overspreading  nearly 
the  entire  under  surface  in  a  large  adult.  About  one  hundred  specimens  se- 
cured" (F.  N.  Blanchard,  Wash.,  1921,  p.  6). 

"Rana  pretiosa  is  found  abundantly  in  the  high  mountain  streams  and  small 
ponds  left  by  the  melting  snow.  It  is  strictly  an  aquatic  frog  and  is  seldom 
found  any  distance  from  water"  (J.  R.  Slevin,  Calif.,  1928,  p.  136). 

"In  the  vicinity  of  Pullman  the  first  amphibian  to  appear  in  the  spring  is 
the  long  toed  salamander,  Ambystotna  macrodactylitm.  .  .  .  The  spotted  frog 
emerges  a  few  weeks  later,  usually  late  February  or  early  March.  By  the 
middle  of  March  and  early  April  the  breeding  season  is  in  full  swing,  andlmat- 
ing  pairs  as  well  as  eggs  in  all  stages  of  development  can  be  found.  The  breed- 
ing season  extends  into  the  middle  of  April.  After  this  time  it  is  usually  im- 
possible to  find  eggs  or  even  adult  frogs"  (A.  Svihla,  Wash.,  1935,  p.  119). 

"Common  along  the  sloughs  of  the  Willamette  and  Columbia  rivers.  It  is 
mainly  aquatic,  seldom  leaving  the  water"  (S.  G.  Jewett,  Jr.,  Ore.,  1936,  p.  72). 

"It  is  found  in  or  near  pools,  ponds,  slough,  marshes  or  springs"  (K.  Gordon, 
Ore.,  1959,  p.  64). 


Map 


522  HANDBOOK  OF  FROGS  AND  TOADS 

"This  is  a  race  of  the  Pacific  northwest  which  occurs  in  the  Rocky  Moun- 
tains in  Montana  and  Wyoming.  It  is  most  abundant  in  the  higher  mountain 
lakes"  (T.  L.  Rodgets  and  W.  J.  Jellison,  Mont.,  1942,  p.  n). 

"Two  specimens  o£  this  species  were  taken  in  the  grasses  at  the  edge  of  Jorn 
Lake,  where  this  form  apparently  meets  the  following  f R.  p.  hiteiventris]" 
(F.  G.  Evenden,  Jr.,  Ore.,  1945,  p.  252). 

Size:  Adults,  2%-$%  inches.  Males,  44-75  mm.  Females,  46-95  mm.  (These 
measurements  are  of  R.  p.  prctiosa  and  JR.  p.  Ittteiventris.)  We  doubt  if  males 
and  females  under  50  mm.  breed  to  any  great  extent.  We  have  seen  records 
of  100  mm.  or  more  for  these  frogs,  but  such  large  specimens  must  be  rare. 
In  Dr.  Storer's  material  we  saw  females  from  46-^0  mm.,  males  from  48-62 
mm.  In  Mr.  J.  R.  Slater's  collection  we  have  females  39-99  +  mm.  and  males 
33-68  mm.,  but  one  36  mm.  we  call  young  (very  warty,  probably  not  mature, 
going  to  be  a  male).  A  male  of  44  mm.  we  think  perhaps  mature,  thumb 
enlarged.  We  did  not  determine  the  sex  of  a  University  of  Michigan  Zoologi- 
cal specimen  of  44  mm.  Specimens  below  44  mm.  are  doubtless  immature. 

General  appearance:  This  is  a  meclmm-si/ed  frog,  light  or  dark  brown  with 
finely  roughened  skin  on  the  dorsal  parts;  the  back  and  top  of  head  have 
inky  black  spots,  which  at  times  may  be  very  large  and  irregular.  Occasionally 
these  dark  spots  have  light  centers  like  the  ones  in  R.  a.  draytonii.  It  usually 
has  a  prominent  dorsolateral  fold,  but  in  some  even  this  disappears.  Some- 
times the  throat  and  the  entire  under  parts  arc  spotted.  The  under  parts  are 
yellowish  to  salmon  reel,  clouded  or  marbled  with  gray.  The  limbs  have  cross- 
bars. Preserved  specimens  may  approach  the  appearance  of  R.  s.  cantabngensis, 
or  with  few  or  no  spots  may  suggest  the  green  frog,  R.  clanntans.  Groin  not 
mottled  as  in  R.  a.  aurora. 

"The  Western  Spotted  fiog  can  be  recogni/cd  by  its  stout  form,  rough  skin 
and  by  the  presence  of  dark  spots  with  light  centers  on  the  back  and  salmon 
or  red  colour  on  the  under  parts  only"  (Ci.  C.  Carl,  B.C.,  1943,  p.  50). 

Color:  Near  west  edge  of  Provo  at  springy  area.  Female,  The  back  is  ser- 
pentine green  or  Saccardo's  olive,  becoming  on  upper  part  of  fore  limbs  and 
sides  yellowish  citrine  or  olive  lake.  The  dorsum  of  this  individual  bears  no 
dark  spots  and  has  no  costal  folds,  so  that  it  reminds  one  of  a  young  bullfrog. 
On  the  sides  are  a  few  flat  tubercles  with  chamois  or  pinkish  cinnamon  tops. 
There  is  a  fold  of  the  same  color  above  the  arm  insertion  that  passes  under 
the  tympanum  but  there  becomes  very  faint.  The  groin  is  somewhat  ihottled. 
There  arc  three  crossbars  of  olive  on  femur  and  tibia,  two  on  tarsus,  and  one  on 
foot.  The  eye  is  almost  black  with  iust  a  slight  touch  of  citron  yellow  spots 
and  a  very  thin  pupil  rim  of  the  same  color.  The  underside  of  throat  is  cream- 
colored  as  is  also  the  venter.  Underside  of  fore  limbs,  forward  half  of  femur, 
underside  of  tibia,  and  top  of  tarsus  onto  inner  side  of  foot  are  zinc  orange, 
apricot  orange,  or  orange  chrome  in  solid  color.  The  pectoral  region  and  belly 
are  thickly  spotted  with  the  same. 


FROGS:  RANIDAE 


523 


Male.  The  dorsum  is  yellowish  citrine  or  olive  lake,  the  same  on  fore  limbs 
and  top  of  head.  The  back  bears  numerous  inky  black  spots  with  a  few  of 
the  rear  ones  light-centered.  The  barring  of  the  legs  is  like  that  of  female. 
There  is  not  quite  so  much  zinc  orange  on  the  belly,  but  on  the  legs  it  is  more 
intense  becoming  scarlet.  The  underside  of  fore  limbs  and  hind  limbs  is  scarlet, 
and  there  is  spotting  of  same  color  on  breast  and  belly.  The  throat  is  cream. 
The  eye  is  like  the  female's.  The  dorsal  fold  is  practically  absent.  Another 
male  is  almost  uniform  warbler  green  or  serpentine  green  on  the  back,  al- 
most like  a  bullfrog.  These  two  males  are  from  Wilmer  W.  Tanner. 

British  Columbia.  "Specimens  were  taken  in  small  streams  near  Watson 
Lake,  Meadow  Lake,  and  Clinton.  Following  is  a  description  of  a  2.75  inch 
female  specimen  from  Meadow  Lake.  Hind  leg  to  heel  equals  length  of  body 
forward  to  ear.  Skin  everywhere  rough.  Lateral  folds  distinct.  Ground  color 
yellowish-brown  on  head,  back,  and  upper  surface  of  legs.  Irregular  roundish 
spots  on  head,  back  (including  lateral  folds),  and  front  legs  arc  black,  and 
the  majority  have  raised  reddish-brown  centres.  Indistinct  leg  bars  formed 
by  black  specks.  Irregular  black  spot  above  each  eye.  Lateral  folds,  reddish- 
brown.  Vertebral  stripe,  light  yellowish-brown  showing  only  on  pelvis.  Ir- 
regular line  from  eye  across  nostril  to  end  of  muzzle,  black  sprinkled  with 
yellowish-brown.  Reddish-cream  streak  from  muzzle  to  arm,  interrupted  at 
end  of  jaw.  Sides  whitish,  mottled  with  yellowish-brown.  Throat  white, 
mottled  with  salmon-red.  Belly  white,  marked  with  salmon-red  in  shape  of 
thick  U.  Legs  salmon-red  on  lower  surface.  Feet  salmon-red  on  inner  half 
(hidden  when  sitting).  Soles  of  leer,  purplish-brown  with  lighter  tubercles. 
Of  five  specimens  1.15  inch  in  length,  only  one  shows  slight  traces  of  salmon- 
red  on  lower  surfaces  of  femur,  all  other  under  surfaces  being  immaculate 
white"  (C.  L.  Patch,  B.C.,  ic>22,  p.  78). 

Structure:  Male  with  thumbs  enlarged  and  webs  of  hind  feel  tending  to 
become  convex  as  in  Rana  svlvatica;  tip  of  fourth  toe  free;  no  dark  check 
patches;  no  red  on  sides. 

"Female.  Body  thick  and  stout;  head  shoit,  broader  than  long.  Tympanum 
not  two-thirds  the  length  of  the  eye.  Tongue  lar^e.  Palatine  teeth  minute, 
posterior  to  the  inner  narcs.  Skin  leathery,  covered  with  asperities,  except  on 
inner  surfaces,  even  on  the  sole  of  the  foot.  A  depressed  ridge  of  skin  on  each 
side,  none  intermediate;  a  glandular  ridge  along  the  upper  jaw.  Femur  not 
half  the  length  of  the  body;  tibia  about  equal  to  ir,  but  shorter  than  the  hind 
foot.  Terminal  joint  of  longest  toe  free,  next  margined,  and  web  generally 
extending  between  the  tips  of  the  toes  on  one  side,  and  the  last  articulation 
on  the  other.  Shortest  toe  rather  more  than  one-third  the  length  of  the  hind 
foot,  both  measured  from  the  tarsus.  Above  yellowish  brown  with  rounded 
dark  blotches.  Sides  dusky;  dorsal  ridge  lighter;  a  light  line  along  the  pos 
terior  ridge  of  the  upper  jaw.  Faint  indications  of  a  dark  area  about  the 
tympanum;  a  few  spots  about  the  nostrils.  Beneath  yellowish  white,  obsoletely 


524  HANDBOOK  OF  FROGS  AND  TOADS 

marmorated  with  brown.  About  two  and  a  half  inches  long.  Syn.  Rana 
pretiosa  B.  &  G.,  Proc.  Acad.  Nat.  Sc.  Phila.  VI  #8.  Hab.  Washington  Ter- 
ritory" (S.  R  Baird,  Gen.,  [1854!  1856,  p.  62). 

Voice:  The  male  is  without  lateral  vocal  sacs. 

"While  endeavoring  to  escape  from  the  hand,  a  young  specimen  1.45  inch 
in  length  opened  the  mouth  and  emitted  squeaks  resembling  those  made  by  a 
mouse"  (C.  L.  Patch,  B.C.,  1922,  p.  78). 

"Soon  after  the  spotted  frog  makes  its  appearance  in  the  spring,  mating 
commences,  and  the  ponds  are  resonant  with  their  deep  bass  calls.  Their  short 
bass  notes  are  easily  distinguishable  from  the  almost  constant  high  shrill  song 
of  the  tree  frog  and  can  be  heard  for  at  least  a  quarter  of  a  mile.  On  several 
occasions  pairs  of  .spotted  frogs  brought  into  the  laboratory  while  in  amplexus 
were  observed  to  drop  to  the  bottom  of  the  aquarium  in  which  they  were 
kept  and  there  emit  their  call.  Although  the  sound  was  greatly  dampened 
by  the  water,  the  note  could  still  be  heard  distinctly  for  some  distance  within 
the  room.  When  emitting  their  call  notes  only  a  slight  tremor  at  the  sides  of 
the  throat  is  visible.  The  vocal  sacs  are  evidently  poorly  developed  for  they 
do  not  protrude  as  they  do  in  other  ranas"  (A.  Svihla,  Wash.,  1935,  pp.  119- 
120). 

Breeding:  This  species  breeds  from  March  to  May.  The  egg  mass  is  about  a 
pint  in  bulk.  The  egg  is  Viu  inch  (2  mm.),  the  outer  jelly  envelope  %-%  inch 
(10-14  mm.)  (adapted  from  Dickerson,  1906,  p.  219).  The  envelopes  are 
large  and  the  eggs  appear  far  apart.  The  tadpoles,  2^-2%  inches  (56-70  mm.) 
with  tooth  ridges  %,  sometimes  %,  transform  from  June  to  August  at  %-% 
inch  (16-23  mm.).  Like  our  eastern  wood  frog,  many  frogs  of  this  species 
apparently  breed  before  summer,  particularly  in  March  to  April. 

"A  female  of  61  mm.  taken  at  Cotton  wood  Creek  on  June  6,  1928,  had 
rather  large  ovaries  with  enlarging  eggs.  Two  females  of  46  and  74  mm.  taken 
at  Six  Mile  lake  on  June  19,  two  of  58  and  60  mm.  taken  at  Lytton,  July  i 
to  8,  1925,  had  only  minute  eggs  in  their  ovaries.  Two  females  of  89  and  93 
mm.  taken  at  Brent's  lake  on  July  i,  and  four  of  from  73  to  78  mm.  taken  at 
Summerland  on  July  6,  1928,  contained  ripening  eggs.  Another  specimen  of 
51  mm.  taken  at  Summerland  on  July  20, 1928,  had  small  ovaries  with  minute 
eggs.  Dickerson  (1906,  p.  219)  gives  the  month  of  March  as  the  spawning 
season  for  this  frog  at  Puget  Sound.  The  above  limited  data  would  suggest 
that  in  the  interior  country  it  occurs  in  June  and  July"  (E.  B.  S.  Logier,  B.C., 

*932>  P-  P3>- 
Tadpole:  The  National  Museum  has  the  following  material: 

No.  21484:  B.  W.  Evermann  and  C.  Gilbert,  Natches  R.  near  N.  Yakima, 
Wash. — i  tadpole  with  4  legs,  total  length  59  mm.,  body  22  mm.;  i  tadpole — 
66  mm.,  body  24  mm.;  i  tadpole — 68.5  mm.,  body  25.5  mm.,  hind  legs  show- 
ing, rather  far  advanced,  but  with  three  rows  of  teeth  on  upper  jaw. 


FROGS:  RANIDAE 


525 


No.  17615:  B.  W.  Evermann,  Silver  Bow,  Brown's  Gulch,  Mont.,  July  27, 
1891 — i  tadpole  56  mm.,  body  23  mm. 

No.  17616 — r  tadpole  53  mm.,  body  25  mm.  Mouth  parts  possibly  not  nor- 
mal but  teeth  %.  (Also  see  account  by  Helen  Thompson,  1913,  under  R.  p. 
lutetvcntns.) 

Transformation:  Time,  June-August,  the  bulk  of  transformation  appar- 
ently in  July  and  early  August.  Size,  16-2}  mm.  July  19,  1908,  Prairie  Hill, 
Selkirk  Mts.,  British  Columbia,  J.  C.  Bradley  secured  transforming  and  trans- 
formed individuals  from  16  to  22  mm.  Others  in  the  same  lot  of  23,  25, 28  mm. 
were  past  transformation. 

The  National  Museum  has  the  following  significant  records: 

No.  32663-6.  June  i,  1898 — 21,  23,  23.5  mm.  Lake  Bennet,  B.C.  A.  Seal. 

No.  53142.  Aug.  2,  1915 — 20  mm.  Riverside,  Matthews  Co.,  Ore.  E.  A. 
Preble. 

No.  17622-4.  Aug.  8,  1891 — 22,  21,  22  mm.  just  past  transformation.  Canon 
Cr.  Nat.  Park,  Wyo.  B.  W.  Evermann. 

No.  16793.  Aug.  20,  1890 — 23  mm.  Salmon  River,  Idaho.  V.  Bailey  and 
Butcher. 

In  Storer's  collection  are  transformees  18,  18.5,  18.5,  and  21  mm.  In  Slater's 
material  a  scries  of  transformed  specimens  range  from  13  to  18  mm.;  another 
series  17  to  27  mm. 

"Nineteen  tadpoles  were  taken  in  Brent's  Lake  on  July  i,  1928.  They  ranged 
m  length  from  43  to  76  mm.  In  two  of  them,  all  four  limbs  were  functioning; 
in  two  more  the  front  limbs  were  well  developed  and  could  be  seen  folded 
beneath  the  skin;  in  seven  others  the  hind  limbs  were  from  one  quarter  to 
half  grown.  .  .  .  There  would  seem  to  be  little  doubt  that  the  mature  and 
transforming  tadpoles  taken  at  Brent's  lake  on  July  i,  with  an  average  body 
length  for  all  of  21.98  mm.,  and  for  the  eleven  larger  specimens  of  27.63  mm., 
belonged  to  the  brood  of  the  preceding  year  and  had  wintered  over  as  tad- 
poles, and  consequently  were  ready  to  begin  transformation  earlier  in  the 
summer  than  those  nearer  the  coast  (Puget  Sound)  which  go  through  their 
metamorphosis  in  the  same  year  in  which  they  are  hatched"  (E.  B.  S.  Logier, 
B.C.,  1932,  p.  324). 

The  best  account  of  its  breeding  is  given  by  A.  Svihla  (Wash.,  1935,  pp. 
119-122): 

"In  the  vicinity  of  Pullman  the  egg  masses  are  deposited  in  shallow  water 
among  the  grasses  at  the  edges  of  ponds,  although  the  egg  masses  themselves 
are  not  attached  to  blades.  .  .  .  Two  egg  masses  were  measured  and  the  num- 
ber of  eggs  estimated.  One  mass  contained  1500  cc.  and  the  other  1 100  cc.  Since 
each  egg  measures  more  than  i  cc.  the  number  of  eggs  in  these  masses  would 
approximate  1500  and  noo  respectively. 

"Each  egg  including  its  jelly  coats  measures  from  TO  to  15  mm.  in  diameter, 


526  HANDBOOK  OF  FROGS  AND  TOADS 

the  egg  proper  being  only  2  to  2.8  mm.  in  diameter.  There  are  two  jelly  coats 
present,  an  inner  one  measuring  from  5  to  6  mm.  and  an  outer  varying  from 
10  to  15  mm.  .  .  .  The  inner  coat  is  rather  hard  to  see  but  is  apparent  when 
viewed  obliquely  or  when  the  egg  becomes  infected  with  a  fungus  growth 
or  bacterial  growth,  which  is  usually  found  in  the  inner  coat.  The  newly  de- 
posited egg  is  black  in  color  at  the  animal  pole  and  creamy  white  at  the 
vegetal  pole.  The  vegetal  pole  varies  in  size  from  a  mere  point  to  half  of  the 
egg.  The  vitellme  capsule  closely  circumscribes  the  egg. 

".  .  .  At  the  time  of  hatching  the  young  tadpoles  are  soot)  brown  in  color 
and  measure  7.$  to  8.7  mm.  in  length.  The  adhesive  discs,  dextral  anus,  and 
external  gills  are  apparent  at  this  time. 

"The  tadpoles  grow  very  rapidly  for  in  about  thirty  days  the  hind  limb  buds 
appear  and  the  total  length  has  increased  from  about  S  mm.  to  36  mm.  At 
this  time  the  head  and  bodv  arc  15  mm.  in  length  and  the  tail  20  mm.  The 
depth  of  the  body  is  about  in  mm.  and  the  greatest  width  7  mm.  The  tail  is 
1.4  to  1.6  times  the  combined  length  of  head  and  body.  The  eyes  are  dorsal, 
well  up  on  the  head,  and  rather  close  together,  the  iiuci orbital  distance  being 
about  2  mm.  The  external  narrs  arc  situated  almost  midway  between  the  eyes 
and  the  tip  of  the  snout  but  are  slightly  closer  to  the  eyes.  The  interorbital 
distance  is  contained  7.5  times  in  the  length  of  the  head  and  body.  The  anus 
is  dextral  and  spiraculum  simstral,  being  directed  upward  and  backward  as 
in  other  Ramdae.  The  greatest  depth  of  the  tail  (over  fins)  is  contained  2.5 
times  in  the  length,  while  the  greatest  depth  of  the  muscular  portion  is  con- 
tamed  approximate!)  7  times  in  the  length  of  the  tail. 

14 At  the  age  of  the  appearance  of  the  hind  limb  buds  the  mouth  parts  appear 
as  follows:  on  the  upper  lip  there  is  but  a  single  low  of  labial  teeth  which  is 
divided  medially  into  two  verv  short  sets  situated  near  the  ends  ot  the  upper 
jaws.  On  the  lower  labitim  there  are  thiee  long  rous  of  teeth,  the  second  or 
middle  row  the  longest,  the  first  and  third  about  equal  in  length.  A  single  rou 
of  labial  cirri  extends  along  the  sides  ami  bottom  of  the  lips.  .  .  . 

"The  mouth  parts  ol  Rm  />  pretw*a  tadpoles  differ  from  those  of  R.  p.  littei- 
I'cntot*.  In  prctw*d  there  are  four  rows  of  labial  teeth,  one  vcrv  short  upper 
row  and  three  lower  imvs.  In  Ititctvcntns  there  arc  five  rows,  two  long  upper 
rows  atul  three  lower,  as  figured  by  Thompson  ( igi  3).  The  first  lower  row  in 
litteii'cntn*  is  longer  than  the  analogous  one  in  pretioM. 

"By  the  middle  ot  June  the  tadpoles  metamorphose  into  frogs.  At  "this  time 
the  newl\  transformed  frogs  are  oo  mm.  in  total  length  with  a  head  and  bod\ 
length  of  2()  mm.  1  here  is  no  indication  ot  the  reddish  coloration  on  the 
underside  ol  the  thighs  and  abdomen  of  these  \oung  frogs. 

"Since  spotted  frogs  ot  two  very  divergent  sr/es,  one  about  half  the  si/e  of 
the  other,  can  be  found  during  the  springtime,  ir  seems  reasonable  to  con- 
clude that  at  least  two  \cars  arc  required  for  this  frog  to  reach  mature  size" 
(A.  Svihla,  Wash.,  KH>  pp.  120-121). 


FROGS:  RANIDAE 


5*7 


Journal  notes:  March  30,  1942.  Went  on  short  trip  with  Mr.  James  Slater 
to  Sparaway  Lake.  Here  he  finds  R.  pretiosti.  (Sparaway  Lake  is  near  old  type 
locality  of  R.  pretiosa.)  Went  to  head  of  Sparaway  Lake  and  traveled  along 
the  drainage  ditches.  Saw  two  frogs  leap  before  we  got  to  them.  Finally  espied 
one,  but  as  I  tried  to  catch  it,  I  went  in  deep.  We  came  back  later  and  caught 
it.  They  are  sluggish,  slow  frogs. 

April  i.  With  Slater,  Sec  R.  aurora  for  key  characters. 

In  one  conversation  with  Slater  he  remarked: 
"No  mottling  in  groin  R.  p.  prctiosa 

Slight  mottling  in  groin  R.  cascaJae 

Always  mottling  in  groin  R.  a.  auroni" 

May  30,  Utah.  With  W.  W.  Tanner  went  i  mile  west  of  Provo  to  a  spring. 
Here  we  found  one  female  Rana  p.  prctiosa  and  one  )oung.  In  another  spring 
tcx)k  five  young  R.  p.  pretiosa. 

Authorities9  corner: 
F.  G.  Test,  Mont.,  [  1891]  1893,  P-  58- 
J.  Cirmnell,  J.  Dixon,  ).  M.  Linsciale,  Calif.,  1930,  p.  144. 

Nevada  Spotted  Frog 

Rana  ptettosa  luteivintn*  Thompson.  Plate  ("XVII;  Map  ^5. 

Range:  Southeastern  Washington,  eastern  Oregon,  noithcastcrn  California, 
northeastern  Nevada,  and  probably  other  interior  parts,  ("amp  (Calif.,  i<ji7* 
p.  124)  holds  that  it  "seems  to  be  represented  in  the  Museum  of  Vertebrate 
/oology  by  two  specimens  (Nos.  2o<jS,  2u<)(j),  collected  by  J)r.  H.  C.  Bryant 
at  Alturas,  Modoc  Countx,  California."  (Jordon  (Oic.,  i<)W,  P-  fy)  assigns 
specimens  to  it  from  Klamath  Lake,  I  Limey,  Ciook,  (Jr.mt,  Umatilla,  Union, 
Wallowa,  and  Linn  counties  and  Dcchutcs  River,  Ore.  Storer,  1925,  assigns 
Fort  Walla  Walla,  Humpcg  Rills,  and  Huitc  ("reek  specimens  from  eastern 
Washington  to  this  foim. 

Do  we  have  R.  p.  ptetiosa  to  the  Cascades,  a  tongue  of  R.  tascadac  down 
the  Oregon  Sky  Line,  and  R.  p.  Inteiventu*  eastward  of  the  Cascades3  The 
specimens  we  collected  at  Lapmc,  Ore.,  we  interpret  as  R.  p.  Intctventrt*. 

In  order  to  be  satisfied  that  the'sc  Oiegon  frogs  were  like  those  of  the  type 
locahu,  we  sought  frogs  from  the  Owyhee-Snake  River  drainage  of  northern 
Nevada.  We  finally  secured  several  handsome  highly  coloicd  frogs  of  this 
species  from  a  boy,  Larry  Flower,  of  Elko,  Nev.,  who  collected  them  in  Taylor 
Canyon,  Llko  Co.,  Nov.,  July  22,  1945.  These  were  as  flame  scarlet  to  aprieol 
orange  on  the  be'lly  as  were  ours  from  Lapmc. 

Habitat:  Irrigation  ditches,  streams,  ponds,  Kikes. 

"It  is  quite  aquatic,  although  the  stomach  contents,  consisting  of  ants  and 
water  insects,  indicate  that  at  least  parr  of  the  hunting  is  done  on  land.  A  few 
\oung  specimens  were  taken  a  short  distance  from  water  on  the  banks  of  Anne 


528 


HANDBOOK  OF  FROGS  AND  TOADS 


Plate    CXVL    Runa    prctiosa    prctiosa 
>  Males.  5.  Female. 


Plate  CAT//.  Rana  prctiosa  lutewentris 
(X:*4).  1,2.  From  Deeth,  Nev.  3.  Enlarged 
thumb  of  male,  from  Lapine,  Ore.  4,  From 
Lapine,  Ore. 


FROGS:  RANIDAK 


5*) 


Creek,  but  the  adults  were  usually  found  along  the  edges  of  swifth  flowing 
streams  or  with  only  the  head  projecting  from  the  vegetation  of  stagnant 
pools.  When  disturbed,  they  slipped  quietlv  under  the  surface,  but  quickly 
reappeared,  usually  in  the  same  place.  Although  the\  were  almost  as  com- 
mon as  Rana  ptpiens,  the  two  species  were  seldom  found  in  the  same  pond" 
(H.  Thompson,  Nev.,  1913,  pp.  53-55). 

Size:  (See  /?.  p.  pretiosa.)  r'^-31^  inches  (44-^7  mm.). 

General  appearance:  "Color  above  grayish  brown.  ...  In  the  brighter 
colored  specimens  the  lateral  folds  are  lighter  than  the  ground  color.  There  is 
occasionally  a  trace  of  a  dark  cheek  patch,  and  the  spots  on  the  back  may  be 
obscure  or  distinct  and  few  or  many  in  number.  The  blight  color  on  the  ven- 
tral surface  vanes  from  a  faint  yellowish  tinge  on  the  feet  of  the  young  speci- 
mens to  orange  yellow  (141,  156,  ifii,  171)  IKhncksicck  et  Valette,  Code  ties 
Cottlenrs,  Pans  |  in  adults,  and  it  may  be  present  on  the  thighs,  may  extend  in 
a  more  or  less  U-shaped  blotch  on  the  belly,  or  may  cover  the  entire  ventral 
surface  to  the  shoulder  girdle"  (H.  Thompson,  Ncv.,  K>I  },  p.  54). 

Color:  Mule.  Deeth,  Nev.,  May  22,  1942.  Back  and  clorsum  of  hind  limbs 
dark  greenish  olive  or  deep  olive,  fore  limbs  citrine-drab  or  yellowish  olive. 
There  is  band  on  upper  ]aw  of  pale  ochraceous-salmon  from  near  tip  of  snout 
to  arm  insertion,  with  a  touch  of  same  on  lower  jaw;  margins  of  both  jaws 
marbled.  The  spots  of  the  back  are  few  (S)  and  very  indistinct,  with  light 
tubercle  in  center.  This  frog  when  first  caught  showed  no  spots  because  it  was 
so  dark.  There  is  no  dark  mask.  The  tympanum  is  smooth,  smaller  than  eye. 
The  ins  is  black  with  flecks  of  ochraceous-orange  above  and  below  pupil  rim, 
which  is  pale  chalcedony  yellow.  The  very  front  of  eye  bears  a  slight  uvsconuc 
bar  of  Veronese  green. 

Female  Decth,  Ncv.,  May  2|,  1942.  The  back  is  dark  olive,  with  a  tew  light 
tubercle-centered  black  spots  which  arc  not  very  evident.  There  is  a  promi- 
nent bar  on  forward  half  of  brachmm,  but  otherwise  bars  on  fore  limb  are 
practically  lacking,  and  bars  on  hind  limbs  are  obscure.  The  costal  iolds  arc- 
broad  and  rounded  but  quite  conspicuous.  An  avellancous  jaw  stripe  extends 
to  the  arm  insertion.  The  upper  labial  margin  is  fuscous  with  specklmgs  of 
light  green-yellow,  the  lower  fuscous  interrupted  by  the  pale  ohve-bufT  of  the 
throat.  On  the  rear  of  thigh  each  tubercle  is  apricot  yellow.  There  is  apricot 
yellow  on  the  sides  of  belly  and  back  of  arm  insertion  and  a  touch  of  it  on 
pectoral  region.  The  background  of  throat  and  belly  is  pale  olive-buff  to  white 
with  fuscous  black  spotting  on  the  belly  and  fuscous  mottling  on  the  throat. 
The  ins  of  eye  is  black  or  dark  olive  heavily  punctuated  with  bright  green- 
yellow.  The  skin  is  rough. 

Lapine,  Ore.,  April  5,  1942.  Adtdt  male.  The  top  of  head,  face,  and  dorsal 
background  are  Saccardo's  umber  or  snufT  brown  becoming  on  arms  and  legs 
light  brownish  olive.  There  is  heavy  black  spotting  on  the  back  and  spots 
on  top  of  head  and  back  of  head  across  the  neck  in  this  individual.  The  spots 


530  HANDBOOK  OF  FROGS  AND  TOADS 

between  costal  folds  and  below  them  have  light  pinpoint  tubercles  as  centers. 
The  tympanum  is  light  brownish  olive;  the  costal  folds  and  slight  suffusion 
over  shoulder  back  of  tympanum  are  warm  sepia.  The  brachium  has  one  half 
and  one  full  black  crossband,  the  antebrachium  has  two  full  and  two  broken 
bands.  In  the  hind  limbs,  the  front  half  of  femur  bears  three  half  black  bands 
less  prominent  than  ones  on  tibia,  tarsus,  and  toes,  which  are  prominently 
banded.  The  intervening  area  is  grayish  olive.  The  membranes  of  hind  foot 
are  suffused  with  vinaceous-brown  or  purplish  vmaceous.  A  jaw  stripe  starts 
at  snout,  not  prominent  ahead  of  eye,  enlarging  below  eye,  and  continues 
to  insertion  of  arm  as  pale  salmon  color  stripe,  which  then  extends  backward 
as  a  body-colored  fold  of  skin  forming  a  half  moon  back  of  axil.  The  edge  of 
the  lower  jaw  is  marked  with  a  series  of  spots.  The  eyes  stand  up  prominently 
and  arc  black  with  a  light  green-yellow  bar  above  the  pupil  and  a  suffusion 
of  the  same  below.  The  color  on  the  belly  is  flame  scarlet  to  apricot  orange 
with  a  few  cephalic  mid-belly  mottlings  of  apricot  buflf.  This  color  extends 
up  the  sides  and,  mingling  with  dorsal  color,  gives  an  English  red  cast  to  the 
lateral  areas.  This  scarlet  to  orange  is  most  pronounced  in  the  groin,  which 
is  not  mottled.  The  flame  scarlet  to  apricot  orange  appears  also  on  the  under- 
side of  arms  and  forward  half  of  femur.  The  lower  third  of  femur  and  tibia 
is  Brazil  red  washed  out  to  peach  red  on  the  upper  tarsus.  These  oranges  are 
not  solid  but  interlaced  or  mottled  with  light  buff  or  cream  color.  The  mottling 
most  prominent  on  the  ventral  body  color  becomes  dots  on  femur.  The  same 
kuul  ocuus  on  upper  pectoral  and  throat  region,  but  there  the  intense  oranges 
are  replaced  by  light  grayish  olive. 

Young.  Upper  part  citrine-drab  with  no  dark  spots  below  dorsolatcral  folds, 
but  between  folds  arc  about  three  spots,  wide  but  not  regular  and  with  light 
cential  tubercle.  The  venter  is  unmoulcd  cream-colored,  with  slightest  touches 
of  orange  on  belly  and  orange  on  underside  of  limbs,  but  lacking  in  groin. 

Second  young.  Just  a  touch  of  orange  on  fore  limb  and  wash  on  femur, 
which  becomes  bright  on  tibia,  tarsus,  and  foot. 

Structure:  "Form  stout.  Head  broader  than  long;  snout  rounded;  external 
narcs  small  and  round,  nearer  end  of  snout  than  eye;  eye  small;  tympanum 
small,  three-fifths  the  size  of  the  eye;  tongue  large  and  fleshy,  strongly  notched 
behind:  vomcrinc  teeth  extending  back  in  two  oblique  patches  from  the  inner 
edge  of  the  mteinal  narcs.  Skin  roughly  tubercular;  sides,  outer  surface  6f 
leg  and  lower  MI i face  of  foot  covered  with  small  pointed  granules;  lateral  folds 
inconspicuous;  glandular  ridge  along  the  jaw,  interrupted  at  the  angle  of  the 
law  and  at  the  shoulder  by  a  ridge  curving  behind  the  tympanum;  fold  of 
skin  across  the  chest.  Fingers  not  webbed,  first  slightly  longer  than  second, 
palmar  tubercle  indistinct.  Legs  massive,  length  to  heel  equalling  that  of  body 
forward  to  anterior  corner  of  eye;  foot  broadly  webbed;  terminal  joint  of 
fourth  toe  free;  small  inner  metatarsal  tubercle,  no  outer  tubercle"  (H. 
Thompson,  Nev.,  19 13,  pp.  55-54). 


FROGS:  RANIDAE 


53* 


Voice:  Not  on  record. 

Breeding:  Probably  from  March  or  April  onward.  Labial  tooth  rows  of  tad- 
pole 2-3.  Miss  H.  Thompson  described  a  tadpole  collected  July  TO  which  was 
neanng  transformation. 

"One  mass  of  260  cc.  volume  contained  about  2,400  eggs  spaced  one-quarter 
to  three-quarters  of  an  inch  from  each  other.  The  jelly  of  the  eggs  is  not  firm 
thus  giving  the  mass  a  loose,  flowing  appearance  when  agitated. 

"Only  one  clear  transparent  gelatinous  envelope  is  present;  its  outline  is 
fairly  distinct,  hut  merges  slightly  with  the  jelly  of  the  adjacent  eggs.  The 
vitellme  capsule  is  close  to  the  vitellus  and  is  seen  best  with  the  aid  of  a  lens. 
The  vitclh  are  black  at  the  animal  pole  and  tan  at  the  vegetal  pole. 

"Eggs  at  late  cleavage  to  crcscentic  groove  stage  gave  the  following  measure- 
ments: vitellus  1.97  mm.  (range  1.81  to  2.12  mm.);  vitelline  capsule  2.06  mm. 
(range  1.9?  to  2.25  mm.);  envelope  6.35  mm.  (range  5.0  to  7.12  mm.). 

"Affinities:  R.  p.  lutetvcntns  eggs  are  much  smaller  in  the  diameter  of  the 
envelope  than  those  of  R.  p.  pretioui,  which  Dickcrscm  (1906,  p.  219)  gives 
roughly  as  being  one-half  inch  across.  Svihla  (1955,  p.  120)  gives  ten  to  fifteen 
millimeters  as  the  diameter  and  also  notes  that  there  is  an  inner  envelope  five 
to  six  millimeters  m  diameter  in  R.  p.  pretiosa  eggs.  No  such  envelope  could 
be  demonstrated  in  our  eggs  of  R.  p.  luteiventns.  Sizes  of  the  vilelli  are  ap- 
parently the  same"  (R.  L.  Livczcy  and  A.  H.  Wright,  Gen.,  1945,  pp.  704- 


Journal  notes:  July  29,  19^4,  Welcome  Motor  Park,  Wells,  Nev.  Went  up 
one  of  the  streams  that  is  supposed  to  be  one  source  of  Humboldt  River  almost 
to  the  fish  hatchery.  The  water  is  cold.  No  luck.  Returned  to  station  and  found 
Anna  had  one  of  the  pri/es  we  sought.  When  she  first  saw  it,  it  was  silting 
just  at  the  edge  of  the  pond  in  the  water.  When  we  caught  it,  it  was  resting 
on  a  mat  of  algae.  It  ducked  beneath  the  algae  and  slowly  swam  out  and  rested 
on  the  muddy  bottom  with  its  head  in  the  mud.  They  arc  rather  slow.  They 
have  considerable  orange  or  yellow.  When  held  in  hand  they  sometimes  give 
a  very  interesting  squeal  but  not  the  mercy  cry  of  the  bullfrog. 

Easter,  April  5,  1942,  Lapinc,  Ore.  Clear  at  1  1  :oo  A.M.  Frosty  last  night.  Shal- 
low weedy  end  of  irrigation  overflow  ditch  beside  road.  Anna  saw  something 
move  in  the  grassy  vegetation.  She  called  to  me.  Just  then  I  espied  a  frog  egg 
mass  in  ditch  on  opposite  side.  I  came  to  her.  We  walked  along  ditch.  About 
iVy  feet  from  edge  of  water  3  inches  deep  is  a  mass  6  by  ^!/6,  another  3  by  $ 
in  water  4  inches  deep.  Anna  and  I  came  to  a  little  pool  where  there  were 
three  masses.  Here  poised  in  water  was  an  adult  R.  pretiosa.  Tried  to  get  it 
with  net.  Never  should  have  lost  it.  Some  of  these  masses  at  surface  were  8 
by  5  or  6  inches.  Saw  one  mass  in  shallow  water  almost  high  and  dry.  While 
I  was  easing  an  egg  mass  out  of  water,  Anna  espied  a  frog  on  opposite  bank 
among  grassy  tussocks.  Egg  masses  gather  a  lot  of  dirt.  One  was  in  center  of 
ditch.  Fresh  ones  are  quite  bluish. 


532  HANDBOOK  OF  FROGS  AND  TOADS 

In  one  pool  were  three  egg  masses  at  surface.  Here  was  an  adult.  Lost  it. 
Later  came  back  to  area  and  caught  it,  a  beautiful  male.  Color  on  belly  dif- 
ferent from  Washington  (Tacoma)  R.  prettosa.  Has  one  metatarsal  tubercle. 
If  one  loses  one  of  these  frogs,  he  reappears,  like  R.  prettosa  of  Tacoma,  in 
the  same  place  poised  on  water  or  in  weeds.  This  is  flat  land,  open,  partly 
cultivated,  partly  meadows,  and  with  trees. 

May  22,  1942,  Wells.  Have  had  only  two  or  three  nice  days  here.  Snowed 
considerably  last  week.  Went  in  creek  below  Welcome  cabins,  8  miles  west 
of  Wells,  but  no  frogs. 

At  Deeth  are  many  sloughs  more  or  less  tributary  as  overflows  of  Mary's 
River.  One  shallow  pond  only  slightly  connected  with  sloughs  from  river 
had  clumps  of  sedges  slightly  out  from  shore.  As  I  walked  along  shore  saw 
quite  a  stirring  movement  in  one  clump.  Bert  came  over,  placed  the  net, 
tramped  in  the  clump,  and  the  frog  came  out  moving  to  one  side.  Bert  swung 
net  around  and  caught  it.  Saw  another  movement  in  near-by  clump.  Bert 
thought  he  saw  a  larger  frog  but  lost  it.  We  made  several  dips  at  the  spot  but 
had  no  more  luck. 

When  Anna  told  me  a  movement  was  in  a  certain  clump,  I  stomped  it  out 
toward  the  net.  Was  lucky  to  get  a  male  Rana  p.  Intewentris.  Soon  saw  another 
movement  and  for  an  instant  a  bigger  one  than  that  captured  appeared  but 
went  back  into  vegetation.  Never  got  it. 

May  23, 1942,  Elko.  Went  to  Maggie  and  Susie  Creeks  at  Carlin.  Streams  too 
muddy,  Humboldt  River  too  high,  winter  snows  not  yet  gone  long  enough 
to  make  hunting  satisfactory. 

In  afternoon,  late,  we  went  to  Deeth  to  our  favorite  pond.  Anna  for  an  in- 
stant saw  two  little  bubbles  in  a  tussock  of  sedge,  but  when  the  two  bubbles 
went  back  into  the  water  she  knew  it  was  a  frog.  We  never  saw  it.  Later  re- 
turned and  here  was  her  "bubble  frog."  Approached  slowly  and  with  quick 
sweep  of  net  I  had  it,  a  half-grown  jR.  prettosa  htteiventns.  Spring  has  barely 
arrived  and  several  fields  are  yellow  with  the  dandelions  in  bloom. 

On  way  to  Salt  Lake  City  stopped  at  9:30  at  our  Deeth  pond.  Thought  I 
saw  a  movement.  Took  off  my  shoos  and  stockings.  As  I  approached  pond, 
something  leaped  in.  Placed  my  net  with  idea  of  driving  the  frog  in  the  clump 
into  it.  Just  then  saw  a  little  one  in  next  clump  and  he  was  facing  the  shore. 
Caught  it  by  hand,  then  trompcd  the  other  clump  and  didn't  look  in  net  at 
once.  Then  to  my  surprise  here  was  a  small  female  in  my  net.  What  luck! 
Our  only  captures  have  been  from  this  one  small  pond, 

Authorities9  corner: 

A.  Svihla,  Wash.,  1935,  p.  119.  J.  M.  Linsdale,  Nev.,  1940,  p.  212. 

K.  Gordon,  Ore.,  1939,  p.  64.  F.  G.  Evendcn,  Jr.,  Ore.,  1943,  p.  252. 

Dr.  Gordon  and  his  Oregon  group  interpret  R.  p.  prettosa  as  entering  Cas- 
cades from  west  and  R.  p.  Intewentns  from  high  Cascades  eastward  to  north- 
ern Nevada.  In  the  very  early  spring  of  1942  our  student,  Miss  Ruth  Hopson, 


FROGS:  RANIDAE 


533 


at  Eugene,  tried  to  take  us  to  Mackenzie  Pass,  but  snow  stopped  us.  She 
described  from  the  'Three  Sisters"  a  frog  such  as  Evenden  records  on  the 
Oregon  Sky  Line,  immediately  north  of  Mt.  Washington.  Later  Slater  showed 
us  his  R.  cascadae  which  he  collects  at  Breitenbush  area  just  north  of  Mt. 
Jefferson.  These  suggest  a  meeting  in  ranges  of  R.  p.  hiteiventris  and  R.  cas- 
cadae in  northern  Oregon. 

We  are  following  Mrs.  Gaige  and  employ  Dr.  Gordon's  interpretation  that 
the  R.  pretiosa  from  Cascades  in  Oregon  eastward  into  northern  Nevada  is 
R.  p.  hiteiventris. 

Mink  Frog,  Northern  Frog,  Hoosier  Frog,  Rocky  Mountain  Frog 

Ran  a  septentnonahs  Baird.  Plate  CXVIII;  Map  30. 

Range:  Northern  New  England  and  northern  New  York  south  to  Peter- 
boro  (G.  S.  Miller,  Jr.),  west  to  Minnesota,  north  in  Canada  to  Hudson  Bay, 
and  eastward  to  Gaspc  Peninsula,  Prince  Edward  Island,  and  Nova  Scotia. 

Habitat:  This  is  an  aquatic  frog,  found  in  peaty  or  sphagnaccous  lakes  or 
ponds  or  in  inlets  or  outlets  of  such  lakes  or  ponds,  particularly  where  water 
lilies  are  growing. 

Size:  Adults,  i%~3  inches.  Males,  4^-71  mm.  Females,  48-76  mm.  "The 
smallest  mature  female  captured  was  48  mm.  in  length  and  the  smallest  mature 
male  46  mm.  It  seems  probable,  therefore,  that  R.  scptentnonalis  females  ma- 
ture at  the  same  age  as  the  males,  whereas  R.  clamitans  females  mature  at  a 
considerably  larger  size  than  do  males  and  probably  at  least  one  year  later" 
(R.  Grant,  Que.,  1941,  p.  152). 

General  appearance:  This  frog  is  a  small  representative  of  the  bullfrog- 
grcen-frog  group.  The  sides  are  heavily  mottled,  the  rear  of  the  femur  is 
heavily  reticulated.  The  back  is  huffy  or  brownish  olive,  almost  uniform,  or 
mottled  with  large  dark  areas  set  off  by  a  tracery  of  light  lines  around  or 
among  them,  or  spotted  with  widely  separated  spots.  Sometimes  the  for- 
ward part  of  the  back  is  uniform  and  quite  green,  and  the  rear  part  spotted 
(much  like  some  /?.  fisheri).  The  upper  jaw  is  green.  The  legs  arc  spotted 
or  with  a  few  bars.  The  mottling  cm  the  femur  is  suggestive  of  the  R.  virga- 
tipes-R.  grylto  group.  In  young  specimens  the  sides  may  be  speckled  and  the 
throats  mottled.  In  the  largest  males,  the  entire  under  parts  are  maize  yellow. 

"These  frogs  are  variable  in  color  and  pattern,  but  are  distinguished  from 
young  R.  clamitans  by  the  less  prominent  dorsolateral  folds,  smooth  skin, 
more  extensive  webs  and  strong  odor  when  handled"  (R.  Grant,  Que.,  1941, 
p.  152). 

Color:  Male.  On  back,  buffy  olive  or  light  brownish  olive  to  olive,  marked 
with  wavy  lines  or  spots  of  oil  yellow  or  yellowish  citrine.  On  the  middle 
of  the  back  are  two  irregular  rows  of  round  or  elongated  black  spots,  widely 
separated.  Back,  ahead  of  tympanum,  is  without  black  spots  and  with  cossc 


534 


HANDBOOK  OF  FROGS  AND  TOADS 


Plate  CXVUL  Rana  septentriondis.  i. 
Female  (X%).  2.  Male  (X1/*).  3-  Egg 
mass  (X1/^).  4-  Head  of  male  (X1/^)-  5- 
Male  (X^)-  6.  Male  (X%). 


Plate  CXIX.  Rana  sylvatica  syfaatica.  I. 
Female  (X%)  •  2.  Foot  of  male  with  con- 
vex webs  (Xi).  3.  Female  (X%)*  4-  Foot 
of  female  with  concave  webs  (XO-  5-  Egg 
mass,  spherical  in  shape  (X1^)-  6.  Male 


FROGS:  RANIDAE  535 

green  or  lettuce  green  interspersed  with  the  huffy  olive.  This  green  is  a  solid 
color  on  upper  jaw.  Courge  or  yellow-green  in  region  of  tympanum,  which  is 
tawny-olive  to  huffy  brown.  Mixture  of  yellow-green,  black,  and  hazel  along 
sides.  Throat,  and  on  some  of  largest  males  entire  under  parts,  maize  yellow. 

Female.  Clear  Lake,  Adirondack  Lodge,  N.Y.,  from  S.  C.  Bishop,  July  14, 
1923.  Several  seem  more  spotted  on  under  part  of  hind  legs  than  are  males. 
Some  have  spots  on  either  side  of  the  throat,  hazel  of  back  more  prominent 
and  more  hazel  on  sides,  usually  more  spots  on  the  head.  Underneath,  fe- 
males are  white  or  whitish  except  for  the  throat,  which  may  be  pale  chalcedony 
yellow  or  seafoam  yellow. 

Structure:  No  lateral  folds,  or  folds  interrupted;  webs  large,  but  leaving 
one  joint  of  fourth  toe  free;  head  narrow,  snout  pointed;  eyes  close  together; 
tympanum  conspicuous,  in  males  much  larger  than  the  eye. 

"Body  stout,  depressed.  Skin  not  tuberculatecl,  but  uneven.  A  broad  de- 
pressed ridge  on  each  side  of  back;  none  intermediate;  a  branch  of  same 
round  the  tympanum,  meeting  a  thickening  from  the  jaw.  Hand  much  longer 
than  forearm.  Femur  and  tibia  nearly  equal,  about  half  the  length  of  the 
body,  and  less  than  the  hind  foot.  Foot  large;  terminal  joint  of  middle  toe 
free;  that  of  the  others  free  only  on  the  inner  side.  Above  light  greenish  olive, 
vcrmiculated  with  lighter,  and  with  a  few  large  dark  blotches  posteriorly. 
Beneath  yellow,  unblotchctl.  No  line  on  sides  of  the  jaw.  Two  inches  long. 
Hab.  Northern  Minnesota"  (S.  F.  Baircl,  Gen.,  |  ^54 1  1856,  p.  61). 

Voice:  This  species  has  a  chant  d'amonr  which  at  chorus  season  can  be 
heard  one-third  or  one-half  mile  away.  A.  H.  Norton  of  Maine  considered  the 
call  similar  to  that  of  the  green  frog,  but  higher  and  slightly  metallic.  He 
thinks  it  resembles  "closely  the  sound  produced  by  striking  a  long  nail  on 
the  head  with  a  hammer  in  driving  it  into  heavy  timber."  S.  C.  Bishop  de- 
scribed the  call  as  a  "Cut-cut,  with  sometimes  a  'burred*  gh-r-r-r."  J.  H. 
Gamier  calls  it  a  "rapid  squeaking  croak."  The  croaking  sac  is  similar  to  that 
of  Rana  palttstns,  being  the  throat  and  sides  of  head  below  the  corners  of  the 
mouth;  it  is  not  so  well  developed  as  in  the  sphagnum  frog  (R.  virgatipes). 

Breeding:  This  species  breeds  from  June  24  to  July  30  or  possibly  Aug.  16, 
the  crest  coming  in  July.  The  egg  mass  is  a  plinth,  3-5  inches  (75-125  mm.) 
by  2  inches  (50  mm.)  thick.  The  egg  is  brown  to  black  and  buff  to  yellow, 
the  envelope  %  inch  (6-7  mm.),  becoming  *^ir,-%  inch  (8-9  mm.)  as  de- 
velopment proceeds.  The  olive  tadpole  is  large,  4  inches  (99  mm.),  its  tail 
elongate  with  acute  tip.  The  tooth  ridges  are  %.  After  a  period  of  i  year  or 
slightly  more,  the  tadpoles  transform  from  June  24  to  Aug.  30,  the  bulk  in 
July,  at  iM«{-i%  inches  (29-40  mm.). 

"Some  of  the  female  mink  frogs  taken  on  July  20  to  22  had  laid  their  eggs. 
Rana  clamitans  and  R.  septentrionalis  thus  occupy  the  same  habitats  and  breed 
at  the  same  time.  Metamorphosing  mink  frog  tadpoles  were  caught  along 
with  metamorphosing  green  frogs  on  July  22"  (R.  Grant,  Que.,  1941,  p.  152). 


536  HANDBOOK  OF  FROGS  AND  TOADS 

Journal  notes:  Near  Dorset,  Ontario,  from  July  2-14,  1913,  we  heard  at 
night  along  the  shore  of  Otter  Lake  the  peculiar  note  which  is  the  croak  of 
the  mink  frog.  On  the  fifteenth  of  July  at  10  P.M.  we  heard  several  frogs  and 
started  with  flashlights  for  Peat  Lake  where  the  species  was  in  chorus.  The 
air  temperatures  ranged  from  52°  to  55  °F.,  but  the  water  of  Otter  Lake  at  its 
surface  registered  69°F. 

In  this  peaty  lake  with  a  clear  sphagnaceous  border  we  found  several 
R.  septentrionalis.  On  July  16  and  17  we  examined  the  place  closely.  All  along 
the  north  edge  ot  the  lake  were  white  water  lilies,  yellow  spatterdocks,  and 
water  shields.  These  three  made  a  perfect  carpet  on  the  water's  surface.  On 
these  plants  during  the  day  the  frogs  rested.  In  the  outlet  to  Otter  Lake  (Ten 
Mile  Creek  between  Lake  of  Bays  and  Otter  Lake)  we  found  them  common, 
July  24,  on  muddy  buttoms  where  water  lilies  were  abundant.  Another  habitat 
we  discovered  Aug.  31  was  a  beaver  lake  where  Cassandra  and  all  the  associated 
hcathlike  plants  grew.  Finally  in  Fletcher  Lake,  Sept.  i,  we  found  them  in 
the  shallow,  sandy  shores  among  pipeworts  (Enocaitlon  articulatum) . 

On  June  14,  1923,  we  found  most  of  our  frogs  at  Hellgatc  Ponds,  Onekio, 
N.Y.  Here  in  driftwood  pools  below  a  beaver  dam,  we  took  several  R.  clamt- 
tans,  R.  catesbeiana  young,  and  R.  septentrionalis.  Later  took  several  more 
mink  frogs  in  muddy  pools  among  granite  boulders. 

Authorities9  corner: 

A.  G.  Ruthven,  Mich.,  1910,  p.  59.  I.  Hoopes,  Me.,  1938,  pp.  4-6. 

P.  H.  Pope,  Me.,  1915,  pp.  1-2.  L.  R.  Aronson,  N.H.,  1943,  pp.  242-244. 

J.  A.  Weber,  N.Y.,  1928,  p.  no. 

Field  notes  of  Dr.  S.  C.  Bishop,  made  at  Clear  Lake,  Adirondack  Lodge, 
N.Y.:  "July  2,  1923.  Many  Rana  septentnonahs  in  Clear  Lake.  24  taken,  of 
which  23  were  males,  i  juvenile  female,  swimming  and  floating  in  open  water 
3  or  4  rods  from  shore  or  resting  on  pads  of  yellow  cow  lilies — croaking  at  8 130 
A.M.  in  broad  sunlight,  at  midday  and  m  the  evening. 

"July  13,  P.M.  Day  mostly  bright  and  windy.  Found  three  egg  masses  of 
the  mink  frog,  Rana  septentnonalts  Haird,  fastened  to  the  stems  of  the  yellow 
cow  lily.  They  were  attached  about  8,  12,  and  18  inches  below  the  surface, 
in  globular  masses  from  4-5%  m.  in  diameter.  The  lilies  were  about  4-6  rods 
from  shore  in  6-8  feet  of  water.  Took  one  egg  mass.  The  egg  mass  is  slightly 
heavier  than  the  water  and  sinks  slowly  when  detached.  The  individual  tggs 
have  3  envelopes  visible  with  a  hand  lens.  First  closely  surrounds  egg — 2d 
about  5  mm.  diameter — last  (3d)  :?s  inch.  The  upper  half  in  freshly  laid  eggs 
black,  the  lower  half  creamy  white.  Division  of  color  being  about  equal.  Frogs 
croaked  just  at  daybreak  and  during  morning,  rarely  in  afternoon.  .  .  . 

"July  i4th.  Frogs  croaked  all  night,  males  on  lily  pads  in  lake  and  a  few 
immature  females  near  shore.  .  .  .  Found  2  egg  masses  .  .  .  ist  about  16  to 
1 8  in.  below  surface,  the  embryos  already  hatching;  2d  lot,  2  feet  below  sur- 
face, freshly  laid.  .  .  .  During  middle  of  day  many  males  left  the  open  lake 


FROGS:  RANIDAE 


537 


to  hide  along  shore.  The  male  croaking  expands  the  throat  and  sides  of  head 
below  ears.  .  .  .  Eggs  are  apparently  laid  at  night—at  least  we  could  find  no 
females  in  act  of  laying  during  the  day  or  early  night  when  using  a  light, 
nor  in  the  early  morn  at  5:30  A.M. 

"July  15.  Dark  and  rainy.  Only  a  few  frogs  croaking  during  the  night  and 
very  infrequently  in  the  daylight.  Three  egg  masses  found.  One  lot  4^-5  feet 
below  the  surface;  2d  with  embryos  ready  to  leave  egg  mass,  3  feet  below  the 
surface,  was  6  in.  in  diameter,  the  jelly  ready  to  disintegrate.  3d  mass,  18-20 
inches  below  surface.  .  .  . 

"July  15,  P.M.  Two  egg  masses,  one  about  4  feet  below  surface  in  7  feet 
water  was  6  in.  in  diameter  and  fastened  to  two  stems  of  yellow  lily — 
Nymphaea  variegata.  The  mass  (with  well  developed  embryos)  was  longer 
than  wide  with  the  supporting  stems  running  through  one  side.  2d  egg  mass 
was  smaller,  about  3x4  inches  and  3  feet  below  the  surface." 

Dusky  Gopher  Frog,  Dark  Gopher  Frog 

Rana  sevosa  Com  and  Netting.  Map  26. 

Range:  "The  known  range  of  sevosa  extends  along  the  Gulf  coast  from  St. 
Tammany  Parish,  Louisiana,  to  Mobile  County,  Alabama.  .  .  .  The  eastern- 
most station  for  sevosa  is  three  hundred  miles  west  of  the  westernmost  capita 
locality;  the  ranges  of  sevosa  and  areolata  arc  separated  by  about  two  hundred 
miles"  (C.  J.  Gcnn  and  M.  G.  Netting,  La.,  1940,  p.  156). 

Habitat:  Gopher  turtle  and  other  holes. 

Size:  Adults:  "The  largest  male  sevosa  among  22  specimens  was  84  mm. 
long,  and  the  largest  of  29  females  was  92.5"  (C.  J.  Gom  and  M.  G.  Netting, 
La.,  1940,  p.  146).  We  ourselves  have  seen  males  71-85  mm.  and  females  82- 
105  mm. 

General  appearance:  "Of  the  three  species  Rana  sevosa  has  the  darkest 
dorsal  coloration  and  the  least  amount  of  contrast  between  ground  color  and 
dorsal  spots.  Many  of  the  paratypes  are  uniform  black  above,  a  condition  that 
appears  to  be  more  characteristic  of  males  than  of  females.  The  lightest  speci- 
mens have  a  gray  or  brown  ground  color  and  dorsal  spots  that  range  from 
red  brown  to  dark  brown  but  are  not  black.  The  dorsal  color  and  pattern 
are  continued  over  the  folds,  which  are  never  distinctively  colored.  The  pre- 
served specimens  offer  no  indication  that  yellow  was  present  on  any  part  of 
the  body  in  life.  Metachrosis  has  not  been  reported  in  this  species  but  may  be 
expected  to  occur  within  a  narrow  color  range. 

"In  sevosa  the  entire  anterior  half  of  the  lower  surface  is  thickly  covered 
with  spots  and  dusky  markings;  in  males  the  remainder  of  the  lower  sur- 
face, except  for  a  small  pubic  area,  is  spotted,  whereas  in  females  the  central 
lower  thigh  surfaces  and  the  posterior  portion  of  the  belly  are  usually  im- 
maculate" (C.  J.  Coin  and  M.  G.  Netting,  La.,  1940,  pp.  150-152). 


538  HANDBOOK  OF  FROGS  AND  TOADS 

Structure:  "A  dark,  medium-sized  Rana  with  a  very  warty  dorsum  and  a 
heavily  spotted  venter.  Its  dorsal  spots  are  frequently  indiscernible  from  the 
dark  ground  color,  but  when  distinct  they  are  irregular  in  shape  and  are  not 
outlined  with  a  light  color.  Rana  sevosa  can  be  distinguished  from  both  races 
of  areolata  by  its  spotted  venter,  lack  of  circular,  light-bordered  dorsal  spots, 
warty  dorsum  with  broad  dorsolateral  folds,  and  broad  head.  Rana  sevosa 
differs  from  capita  in  having  a  darker  ground  color,  heavier  and  more  ex- 
tensive ventral  markings,  broader  waist,  narrower  dorsolateral  folds,  wartier 
dorsum,  and  dark  hind  limb  bars  which  are  broader  than  the  light  spaces 
between  the  bars"  (same,  p.  138). 

"Head  triangular  in  outline,  dorsolateral  folds  high  and  relatively  narrow; 
dorsum  with  numerous  prominent  warts;  dorsal  spots  poorly  differentiated 
from  gray,  brown,  or  black  ground  color;  venter  always  spotted  at  least  from 
chin  to  midbody;  dark  bars  on  hindlegs  separated  by  interspaces  that  are  never 
wider  than  the  bars"  (same,  p.  146). 

Voice:  Little  of  record.  See  next  paragraph. 

Breeding:  "Fortunately,  while  this  description  was  in  course  of  prepara- 
tion, Stewart  Springer  visited  Pittsburgh  and  contributed  additional  infor- 
mation upon  the  habits  of  sevosa  from  memory.  He  recalled  finding  these 
frogs  breeding  in  the  water  in  southern  Mississippi  concurrently  with  Hyla 
gratiosa  and  Hyla  anerea  ctnerea;  Allen  .  .  .  reports  the  former  breeding 
near  Biloxi  on  April  18  and  19,  and  the  latter  'as  soon  as  the  weather  becomes 
warm  and  settled.1  Mr.  Springer  further  reported  that  the  eggs,  in  masses 
about  the  size  of  two  fists,  are  laid  under  water  at  a  depth  of  approximately 
one  foot  and  are  attached  to  plant  stems.  He  stated  that  sevosa  is  less  restricted 
to  cypress  swamps  for  breeding  sites  than  is  gratiosa,  since  the  former  occurs 
also  in  pine  barren  ponds,  even  those  of  temporary  character.  The  call,  as 
he  remembered  it,  is  less  snore-like  than  is  that  of  capita.  He  found  that 
frightened  individuals  dive  and  swim  along  the  bottom  of  the  pond"  (same, 
pp.  142-143). 

Journal  notes:  Most  of  our  knowledge  of  this  form  comes  from  (join  and 
Netting.  We  revert  to  their  original  designation  /?.  sevosa.  We  have  known 
more  or  less  about  it  since  our  visits  to  R.  areolata  territory  in  Illinois.  In  1950 
we  wrote:  "Rana  aesopus  (capita)  South  Carolina  to  Florida  to  Louisiana 
(North  Slidell,  May  5, 1926,  P.  Viosca,  Jr.,  and  H.  B.  Chase  et  al.)."  When*  we 
visited  Stewart  Springer  and  Morrow  J.  Allen  at  Biloxi  we  saw  their  Rana 
sevosa.  On  June  10,  1930,  we  started  from  New  Orleans,  La.,  for  the  north 
country,  Pearl  River,  and  the  pine  barren  parishes  of  Louisiana  with  Percy 
Viosca,  Jr.,  and  H.  B.  Chase.  They  tell  me  that  they  find  R.  aesopus  (sevosa) 
in  holes  under  stumps.  First  found  it  breeding  in  a  mayhaw  pond.  In  gen- 
eral our  luck  with  this  elusive  form  has  been  failure.  One  has  to  be  in  their 
region  at  certain  periods  for  success. 

We  have  these  two  notes:  (i)  Biloxi,  Miss.,  Caribbean  Biol.  Lab.,  1930. 


FROGS:  RANIDAE 


539 


Males,  71,  85,  87,  90,  92  mm.  Females,  82,  82,  84,  86,  90,  90,  105  mm.  Male  78 
mm.  from  12  miles  south  of  Vestry  near  Harrison-Jackson  County  line,  Miss., 
M.  J.  Allen.  (2)  The  sizes  of  these  frogs  are:  82  mm.  male,  very  rugose,  with 
triangle  from  arm  inward  on  venter  conspicuous;  75  mm.  male,  vocal  sacs 
beginning  to  enlarge;  74  mm.  male,  thumb  enlarged,  vocal  sacs  partially  de- 
veloped. Other  males  7*5,  82, 82,  82, 83, 83,  83.5, 85, 85, 90  mm.  Females,  82  mm., 
spent;  85  mm.,  ripe;  88  mm.,  not  ripe;  89  mm.,  ripe. 

Authorities'  corner:  "Piactically  all  extant  specimens  of  sevosa  were  col- 
lected by  Morrow  J.  Allen,  Stewart  Springer,  or  their  associates,  in  southern 
Mississippi.  Neither  they  nor  any  other  collectors  have  secured  either  capita 
or  areolata,  as  now  restricted,  in  this  area"  (C.  J.  Coin  and  M.  G.  Netting,  La., 
1940,  p.  142). 

The  observations  of  Morrow  J.  Allen  (Miss.  19323,  p.  9)  are:  "This  species 
has  been  abundantly  found  throughout  the  months  of  October,  November 
and  December  in  the  burrows  made  by  Gopherus  polyhemits.  When  the  tem- 
perature rises,  these  frogs  become  active  and  may  be  seen  sitting  in  the  open- 
ings of  the  tunnels  down  which  they  disappear  at  the  least  indication  of  dan- 
ger. In  colder  weather  they  are  never  at  the  surface  and  can  only  be  taken  by 
digging  to  the  bottom  of  the  gopher  hole,  where  never  more  than  one  is  found 
in  company  of  one  or  two  turtles." 

"At  the  time  of  publication  of  our  description  of  Rana  sevosa  (1940,  Ann. 
Carnegie  Mus.  28:  137-168)  we  had  not  seen  any  Alabama  specimens  of  this 
species,  but  we  referred  Loding's  record  (1922,  Alabama  Mus.  Nat.  Hist., 
paper  no.  5:20)  of  Rana  areolata  from  Dog  River,  Mobile  County  to  the 
synonymy  of  sevosa  on  grounds  of  geographic  probability.  The  apparent  cor- 
rectness of  this  action  was  confirmed  by  Mr.  Lciding,  who  wrote  under  date 
of  February  12,  1941,  as  follows: 

"  'There  can  be  no  doubt  that  the  Rana  sevosa  described  is  the  same  thing 
as  the  three  frogs  I  took  years  ago  under  drift  logs  on  the  beach  of  Mobile 
Bay  just  south  of  the  mouth  of  Dog  River,  and  which  I  identified  as  areolata 
from  Dickerson's  Frog  Book;  later  Hurtcr  agreed  that  while  it  differed  in 
some  respects  from  areolata  as  he  knew  it  from  Missouri  it  must  be  that  species. 
The  triangular  head,  dorsal  warts,  and  color  agree  entirely  with  your  descrip- 
tion and  the  plate  is  a  dead  ringer  of  these  specimens.  This  frog  must  be  very 
secretive  for  I  have  never  seen  specimens  since,  but  Viosca  identified  the  sound 
of  what  he  called  aesoptis  from  a  swamp  in  the  same  neighborhood.  Two  of 
the  specimens  in  the  Chas.  Mohr  Museum  were  lost  some  years  after,  but  if 
I  remember  right,  Hurter  got  the  third  one.' 

"Mr.  R.  W.  McFarland,  of  Fairhope,  Alabama,  recently  sent  us,  for  de- 
termination, a  gopher  frog  (Alabama  Museum  of  Herpetology  No.  218), 
which  he  collected  on  the  evening  of  June  2,  1941,  as  it  was  hopping  across  a 
road,  8  miles  southeast  of  Fairhope,  Baldwin  County,  Alabama.  This  speci- 
men is  clearly  referable  to  sevosa,  and  thus  constitutes  the  first  record  of  the 


540  HANDBOOK  OF  FROGS  AND  TOADS 

species  east  of  Mobile  Bay.  It  is  an  adult  female,  99  mm.  in  snout-to-vent 
length,  and  the  largest  known  sevosa:  The  maximum  size  of  29  Mississippi 
females  previously  reported,  was  92.5  mm.  It  agrees  with  typical  sevosa  in 
morphology  and  in  ventral  markings,  but  differs  somewhat  in  dorsal  pat- 
tern; the  ground  color  is  lighter  gray,  and  the  dark  spots  superimposed  upon 
it  are  somewhat  larger  and  less  numerous  than  in  most  sevosa.  This  variation 
probably  indicates  only  that  sevosa  is  variable  in  number  and  size  of  dorsal 
spots,  as  are  other  representatives  of  the  areolata  group.  It  is  possible  that 
the  atypical  dorsal  pattern  of  this  specimen  may  indicate  some  capito  tenden- 
cies jn  the  population  east  of  Mobile  Bay,  and  that  sevosa  and  capito  may  be 
found  to  intergrade  somewhere  in  the  area  between  Baldwin  County,  Ala- 
bama, and  Berrien  County,  Georgia,  although  no  gopher  frogs  are  as  yet 
known  from  that  region. 

"Upon  the  basis  of  its  structure  and  relationships  we  postulated  (op.  cit.: 
154)  that  sevosa  was  both  batrachophagous  and  insectivorous.  The  latter  of 
these  suppositions  has  now  been  confirmed,  for  Mr.  McFarland's  specimen 
contained  the  remains  of  three  large  beetles;  namely,  a  carabid  of  the  genus 
Pasimachus,  and  two  scarabaeids  belonging  to  the  genera  Canthon  and 
Ligryus. 

"At  the  Gainesville  meeting  of  the  Society,  Mr.  Percy  Viosca,  Jr.,  exhibited 
two  sevosa,  an  adult  collected  at  Pearl  River,  St.  Tammany  Parish,  Louisiana, 
on  December  29,  1956,  and  a  juvenile  collected  'ten  miles  out  of  Picayune, 
Mississippi*  (Pearl  River  Co.?)  in  August,  1955.  These  specimens  provide 
two  additional  localities  for  the  species"  (M.  G.  Netting  and  C.  J.  Goin,  Ala., 
1942,  p.  259). 

Wood  Frog,  Woods  Frog 

Rana  sylvatica  sylvatica  Le  Conte.  Plate  CXIX;  Map  36. 

Range:  From  southern  Minnesota  and  Wisconsin  south  to  Arkansas, 
Tennessee,  and  northern  South  Carolina,  through  Appalachians  to  Maryland, 
north  to  Nova  Scotia,  thence  westward  through  Quebec  to  Sault  Sainte  Marie. 

Habitat:  Wooded  areas.  Breeds  in  leaf-laden  ponds  and  transient  pools  of 
wooded  districts.  Hibernates  in  logs,  stumps,  under  stones  in  wooded  ravines, 
or  beneath  boards  near  woods,  never  in  the  water. 

Size:  Adults,  rKr^1/^  inches.  Males,  34-60  mm.  Females,  34-68  mm.  One 
female  from  Linville,  N.C.  (USNM  no.  55159),  measures  82.5  mm. 

General  appearance:  The  wood  frog  is  medium  in  size,  either  light  or 
reddish  brown  above,  with  a  darker  brown  streak  or  mask  on  either  side 
of  the  head  and  a  dark  line  from  the  eye  to  the  tip  of  the  snout.  There  is  a 
light  line  along  the  upper  jaw  continuing  to  the  shoulder.  Underneath,  it  is 
a  glistening  white  with  a  dark  bar  on  the  upper  arm.  The  legs  are  long, 
marked  with  dark  crossbars.  It  has  prominent  dorsolateral  folds.  When  caught 


FROGS:  RANIDAE 


541 


in  the  breeding  pools,  the  body  seems  broad,  flat,  rather  soft,  and  dark  in 
color,  but  when  caught  later  in  the  woods,  the  form  is  compact,  more  slender, 
and  the  frog  has  a  very  alert  appearance.  Some  individuals  have  inky  dashes 
on  the  sides  and  occasionally  on  the  back. 

Color:  Male.  Girl  Scout  Camp,  Ithaca,  N.Y.,  Aug.  21,  1929.  The  dorso- 
lateral  folds  are  buff-pink  to  light  vinaceous-cinnamon  or  vinaceous-cinnamon. 


Ran  a  s.  sylvatica 


Map  36 

The  upper  back  may  be  onion-skin  pink  or  vmaceous-fawn.  This  becomes 
on  top  of  head  and  in  front  of  eye  vinaceous-buff.  The  face  mask  and  the 
line  from  eye  to  nostril  are  olive-brown.  The  face  mask  in  the  rear  is  bordered 
by  clove  brown.  The  rear  of  the  femur,  tibia,  and  top  of  foot  are  russet- 
vinaceous  or  vinaceous-russet.  The  dark  bar  on  the  hind  foot  and  on  to  the 
tip  of  the  fifth  toe  is  black.  The  dorsolateral  fold  has  a  few  spots  of  black  or 
clove  brown.  Across  the  arm  insertion  on  the  ventral  side  is  a  black  or  clove 
brown  bar.  The  lower  jaw  has  bars  of  army  brown  or  russet-vinaceous  or 
vinaceous-russet.  The  upper  jaw  margin  is  a  more  or  less  entire  line  of  the 
same  color  with  a  very  prominent  line  above  it,  from  the  nostril  to  below  the 
rear  of  the  face  mask,  which  is  cartridge  buff  to  pale  cinnamon-pink.  The 


542  HANDBOOK  OF  FROGS  AND  TOADS 

under  parts  are  white.  The  rear  half  of  under  parts  of  femur  may  have  some 
pale  orange-yellow  or  buffy  yellow,  soon  yielding  to  the  rear  color  of  the 
femur,  tibia,  and  upper  side  of  foot.  Upper  portion  of  iris  is  vinaceous- 
cinnamon.  Band  through  center  eye  is  black  or  color  of  preorbital  stripe. 
Pupil  rim  citron  yellow,  broken  below. 

Structure:  Head  pointed;  legs  long;  webs  of  hind  feet  of  males  very  con- 
vex at  breeding  season;  hind  leg  long  (0.53-0.62  in  length  of  body  while 
0.62-0.74  in  R.  f.  cantabrigensis).  Tibia  more  than  half  the  length  of  the  body 
(1.67-1.88  in  the  length  of  body). 

"Colour  above  .  .  .  [varies]  from  light  to  dark  brown,  with  two  inter- 
rupted longitudinal  lines  of  black,  a  dark  brown  stripe  extending  from  the 
tip  of  the  nose  through  the  eyes,  and  covering  the  auricles"  (J.  Le  Conte,  Gen., 
1825,  p.  282). 

See  K.  P.  Schmidt's  suggestive  article  (Gen.,  T938d),  "A  Geographic  Varia- 
tion Gradient  in  Frogs,"  for  a  discussion  of  wood  frogs. 

Voice:  The  wood  frog's  note  is  very  short,  a  sharp,  snappy  clac^  At  times 
2,  4,  or  6  notes  are  given  in  rapid  succession;  and  when  close  at  hand,  they 
sound  high  and  grating  in  character.  It  can  be  heard  only  a  short  distance 
from  the  pond.  In  chorus,  it  is  more  of  a  rattle  than  that  of  any  other  frogs. 
It  has  often  been  likened  to  the  quacking  of  ducks. 

"The  note  of  sylvatica  peculiar  to  the  mating  season  I  have  only  seen  or 
known  to  be  given  by  the  males,  and  only  when  in  the  water;  it  consists  of 
a  repeated  croak  or  quack,  and  may  be  heard  at  any  time  of  day  or  night 
when  the  temperature  is  favorable.  The  mouth  remains  closed  and  each  time 
the  sound  is  uttered  the  side  of  the  body  over  each  lung  is  distended  into  an 
oblong  sac.  This  clamor  has  often  been  mistaken  for  ducks  quacking,  but 
when  near  enough  one  finds  it  more  liquid  in  tone.  In  'Early  Spring  in 
Massachusetts'  p.  228,  Thoreau  describes  the  note,  'Wurrk  wurrk  wurr  r  k 
wurk.'  During  the  mating  season  the  males  will  give  voice  in  confinement, 
under  favorable  conditions  of  water  and  temperature.  In  addition  to  the  notes 
already  described,  I  have  heard  the  young  frogs  reared  in  captivity  give  a 
single  musical  'chip,'  not  unlike  the  call  note  of  the  song-sparrow.  The  note 
was  uttered  at  intervals,  in  the  excitement  of  capturing  flies  on  which  they 
were  fed.  Owing  to  the  frog's  extreme  shyness  I  have  never  been  able  to  de- 
tect the  sound  in  the  field"  (M.  H.  Hinckley,  Mass.,  1884,  p.  86). 

Breeding:  This  species  breeds  from  March  19  to  April  30  (Ithaca,  N.Y.), 
at  a  temperature  of  about  5O°F.  The  egg  mass  is  globose,  2/^-4  inches  (62- 
TOO  mm.)  in  diameter,  the  individual  eggs  appearing  larger  than  in  R.  pipiens 
and  freer,  and  the  outer  envelope  of  each  egg  keeping  its  spherical  form.  The 
egg  is  M.4-Vio  inch  (1.8-2.4  mm.),  the  inner  envelope  faint,  /4-/4  inch  (3.6- 
5.8  mm.),  the  outer,  distinct  %-%  inch  (5.2-9.4  mm.).  The  egg  complement 
is  2,000-3,000.  The  deep  olive  tadpole  is  medium,  2  inches  (49.8  mm.),  its  tail 
long,  acuminate,  and  with  dorsal  crest  very  high.  The  tooth  ridges  are  %. 


FROGS:  RANIDAE  543 

After  44  to  85  days,  the  tadpoles  transform  from  June  8  to  Aug.  i,  mostly  be- 
fore July  15,  at  %-%  inch  (16-18  mm.). 

"The  eggs  are  deposited  near  the  shore  where  the  frogs  first  congregate  and 
the  temperature  of  the  water  is  most  favorable.  They  are  of  a  deep  chocolate- 
brown  color  on  the  upper  surface,  and  whitish  beneath.  When  first  laid 
they  are  held  closely  together  by  a  gelatinous  substance,  which,  after  a  few 
hours  in  the  water,  begins  to  increase  in  size,  and  gradually  changes  from  a 
bluish  to  greenish  tint.  These  masses  of  eggs  are  attached  to  grasses,  weed 
stalks,  or  branches  under  water.  It  is  not  uncommon  to  find  two  or  more 
bunches  laid  closely  together,  giving,  as  soon  as  the  gelatinous  portion  en- 
larges, the  appearance  of  one  large  bunch.  This  season  I  found  in  a  swamp 
within  a  space  of  about  ten  by  eighteen  inches,  sixteen  bunches,  evidently  laid 
about  the  same  time,  attached  to  the  reclining  grasses  of  a  submerged  tussock. 
The  masses  of  eggs  vary  little  in  size.  On  counting  a  bunch  of  average  size 
the  day  after  it  was  laid,  which  measured  about  four  inches  long  by  three 
wide,  I  found  it  numbered  1380  eggs,  each  of  which  was  enclosed  by  two 
transparent,  membranous  shells.  The  time  in  which  the  eggs  develop  de- 
pends chiefly  on  the  temperature  of  the  water;  those  laid  early  in  March  are 
rarely  hatched  before  the  first  week  in  April  here,  while  those  laid  at  the 
latter  date,  in  an  average  season,  hatch  in  from  ten  to  fourteen  days.  This 
year,  eggs  laid  March  23,  and  April  2,  hatched  at  the  same  time,  April  16.  In 
this  locality  the  period  of  egg-laying  is  usually  over  about  the  twentieth  of 
April"  (M.  H.  Hmckley,  Mass.,  1884,  pp.  86-87). 

L.  W.  Wilson  (W.  Va.,  1946,  pp.  39-41)  found  these  frogs  breeding  as 
early  as  Feb.  20  in  West  Virginia. 

Journal  notes:  Southeast  Slaughter  House  Pond,  Ithaca,  N.Y.,  April  6,  1908. 
As  I  approached  the  pond  I  heard  the  greatest  chorus  of  low  grating  croaks 
I  ever  recorded  from  wood  frogs.  This  was  at  11:30  A.M.  From  the  south  I 
crawled  upon  them  (belly wise  using  elbows  to  move  along  slowly).  At  times 
used  my  opera  glasses.  In  this  way  was  able  to  reach  the  edge  of  pond  without 
disturbing  croakers.  In  this  pond,  there  were  at  least  200  males  croaking.  It 
resembled  a  small  toad  concourse.  The  same  scrabbling  and  zeal  of  mating. 
The  water  temperature  was  58°.  Where  yesterday  there  was  only  one  Rana 
sylvatica  bunch,  there  are  now  17  bunches  of  eggs  laid  April  5-6.  When  I 
arose,  the  frogs  all  disappeared  simultaneously,  and  to  go  through  the  pond 
one  would  little  realize  200  males  were  there,  to  say  nothing  of  the  females. 
One  pair  was  captured.  One  male  taken.  Near  the  sedge  with  17  bunches, 
found  another  pair.  There  were  about  15  eggs  about  the  vent  of  the  female. 
Was  she  laying  on  the  sedge  when  I  rushed  into  the  pond? 

Since  many  of  the  egg  masses  are  laid  near  the  edges  of  shallow  ponds, 
many  egg  masses  are  left  high  and  dry.  A  far  more  serious  source  of  danger 
is  freezing.  Most  of  the  bunches  are  laid  from  %  to  2  inches  beneath  the 
water's  surface.  In  many  instances,  long  before  the  hatching  period  ap- 


544  HANDBOOK  OF  FROGS  AND  TOADS 

preaches,  the  tops  of  these  complements  appear  at  the  surface  because  of  the 
reduction  of  water  level  by  rapid  evaporation.  In  1907  on  April  n  the  tops 
of  exposed  masses  in  two  ponds  were  frozen  in  a  crust  of  ice  and  spoiled.  The 
following  year,  on  April  17,  in  another  pond,  a  thin  transparent  crust  formed, 
which  was  sufficient  to  kill  the  upper  part  of  several  bunches.  In  all  cases 
observed,  however,  the  lower  portions  not  caught  within  the  sheet  of  ice 
have  progressed  nicely. 

This  silent  species  is  not  often  recorded  in  autumn.  Our  late  records  are 
mostly  in  the  month  of  October,  the  latest  being  Oct.  30,  1905.  Through  mid- 
summer (including  the  first  weeks  of  September)  one  occasionally  comes 
upon  this  form  in  the  woods  in  much  lighter  livery  than  in  the  early  spring. 
Thereafter,  it  begins  to  think  of  its  winter  quarters,  and  by  the  last  of  Sep- 
tember or  the  first  of  October  it  seeks  cover. 

Authorities'  corner: 
M.  H.  Hinckley,  Mass.,  1884,  pp.  87-88. 
A.  M.  Banta,  Gen.,  1914,  p.  172. 

Northern  Wood  Frog,  Cambridge  Frog 

Ran  a  sylvattca  cantabngensts  Baird.  Plates  CXX;  CXXI;  Map  36. 

Range:  Alaska  and  Mackenzie  across  northern  British  Columbia,  Alberta, 
and  Saskatchewan  to  Red  River  of  the  North,  northern  Minnesota,  Wisconsin, 
and  Michigan.  From  Sank  Sainte  Mane  eastward  between  Hudson  Bay  and 
Great  Lakes  drainages  to  Saguenay  River  and  to  Gaspe  Peninsula,  and  into 
northeastern  Maine.  Northern  border  from  Mackenzie  and  Alaska  to  Hudson 
Bay  to  Labrador. 

We  are  not  believers  in  three  forms  (sylvatica,  cantabngensis,  and  latiremis). 
Rather  are  we  close  to  the  Patch  school,  although  we  have  drawn  the  southern 
border  of  R.  s.  cantabngensts  considerably  south  of  where  Patch  ostensibly 
would  place  it.  He  quite  likely  is  right. 

The  paper  by  R.  H.  Howe,  Jr.,  on  North  American  wood  frogs  (Gen., 
1899)  upon  which  we  were  brought  up  says  that  the  so-called  Cambridge  frog 
type  "has  been  lost  to  all  intents  and  purposes,  although  it  may  be  an  old, 
shrunken  and  unlabeled  specimen  found  with  a  lot  from  Saskatchewan,  Can., 
in  the  collection  of  the  Museum  of  Comparative  Zoology."  Howe  recognized 
three  forms  R.  sylvatica,  R.  cantabrigensts,  and  /?.  c.  lattremis.  He  was  helped 
by  S.  Garman,  G.  M.  Allen,  and  F.  W.  True. 

Habitat:  Terrestrial  except  at  breeding  periods,  living  in  woods  among  dead 
leaves  or  moss. 

Size:  Adults,  i%-2%  inches.  Males,  36-50  mm.  Females,  37-56  mm, 

General  appearance:  This  is  a  short-legged  northern  form  of  the  wood  frog, 
with  a  dark  vitta  or  mask  from  the  eye  over  the  tympanum,  and  with  a  light 
line  along  the  jaw.  Many  of  these  small  wood  frogs  are  grayish  in  color  with  an 


FROGS:  RANIDAE 


545 


Plate  CXX.  Rana  sylvatica  cantabrigensis 
(X%).  i,4.5.  Females.  2,3.  Males. 


Plate  CXXL  Rana  sylvatica  cantabrigcn- 
sis.  Adults,  from  Clinton,  B.C.  Negatives  of 
National  Museum,  Canada.  Kindness  of 
C.  L.  Patch. 


546  HANDBOOK  OF  FROGS  AND  TOADS 

irregular  dark  brownish  band  on  the  back  and  a  light  median  dorsal  stripe. 
In  some,  this  stripe  extends  from  snout  to  vent;  in  others,  to  varied  distances 
or  is  entirely  lacking.  There  is  a  prominent  dark  brown  stripe  on  the  snout 
joining  the  dark  mask  back  of  the  eye.  This  is  bordered  below  by  a  light 
yellowish  or  buffy  stnpe.  The  costal  fold  is  bronzy.  In  many  frogs  there  may  be 
dark  spots  bordering  the  costal  fold  below,  on  the  dorsum,  or  on  the  sides. 
Some  are  almost  free  of  dark  spots.  The  throat  and  breast  are  usually  speckled. 
We  are  deeply  grateful  for  a  beautiful  series  of  wood  frog  photographs  from 
Clinton,  British  Columbia.  The  prints  were  loaned  to  us  by  Clyde  S.  Patch 
of  the  National  Museum  of  Canada,  which  kindly  permits  their  use  in  our 
second  plate  for  this  form. 

Color:  The  majority  of  wood  frogs  from  Pembina,  N.D.,  seem  to  possess 
the  light  median  dorsal  stripe,  although  it  is  by  no  means  universal.  Those  of 
Itasca  Park,  Bemidji,  and  points  further  south  are  more  inclined  to  brown 
without  the  light  median  stripe;  the  hind  legs  average  shorter  than  do  those 
of  the  typical  R.  s.  sylvattca;  and  the  frogs  attain  breeding  characters  at  a 
much  smaller  size  than  do  R.  s.  sylvattca. 

From  examination  of  material  in  the  National  and  other  museums,  we 
observed  that,  sometimes  in  the  same  lot,  there  may  be  individuals  of 
the  same  sex,  one  with  middorsal  line,  one  without  it.  In  the  same  way, 
the  light  tibial  stripe  may  be  absent  or  present  in  the  same  lot. 

Pembina,  N.D.,  August,  1930.  Male.  This  frog  has  much  the  same  color  as 
the  female,  but  the  median  dorsal  stripe  extends  only  from  snout  to  point 
halfway  to  vent.  The  costal  folds  are  more  evenly  bordered  below  by  mummy 
brown  spots.  In  the  same  way,  the  dark  grayish  olive  dorsal  band  is  at  times 
edged  with  or  dotted  with  mummy  brown  spots.  The  eye  is  much  the  same 
as  in  the  female. 

Female.  The  median  stripe  extends  all  the  way  down  the  back  from  tip  of 
snout  to  vent.  It  is  tilleul  buff  in  color.  There  is  a  broader  band  of  darker  color 
on  either  side  of  this  light  stripe  that  is  drab  or  grayish  olive.  Between  this 
area  and  costal  fold  is  a  pale  smoke  gray  stripe  extending  backward  from 
the  upper  eyelid.  The  costal  fold  is  avellaneous.  The  costal  fold  is  interrupted 
in  the  rear  but  continued  to  the  hind  leg.  Below  the  costal  fold,  just  back  of 
the  arm  insertion  and  above  the  groin,  are  two  large  mummy  brown  spots. 
There  are  a  few  scattered  spots  of  the  same  color  on  the  sides.  The  mask  is 
mummy  brown,  bordered  below  by  an  ivory  yellow  or  seafoam  green  line, 
which  extends  to  the  snout.  The  margin  of  the  upper  jaw  is  buffy  olive.  The 
lower  jaw  has  a  series  of  prominent  buffy  olive  spots  interrupted  by  seafoam 
green  and  pale  cinnamon-pink  of  the  chin  color.  This  same  green  color  is  on 
entire  venter  of  frog.  The  breast  and  throat  are  spotted  with  an  indefinable 
gray.  The  under  surfaces  of  hind  legs  are  russct-vinaceous.  The  hind  legs  have 
3  or  4  transverse  bars  on  front  half  of  femur;  4  or  5  on  tibia,  about  3  on  tarsus, 
and  2  or  3  on  foot.  There  is  mignonette  green  or  deep  chrysolite  green  in  groin. 


FROGS:  RANIDAE 


547 


The  eye  is  the  same  color  as  the  mask  except  for  the  upper  pupil  rim,  which 
is  strontian  yellow  with  an  ochraceous-orange  band  above  it.  There  is  a  black 
band  on  front  of  brachium  near  arm  insertion. 

Male.  Manitowish,  Vitas  Co.,  Wis.,  Oct.  12,  1929,  from  Carolyn  Wcbcr. 
Back  sayal  brown,  on  sides  cacao  brown.  Lateral  fold,  fore  limbs,  and  some 
of  hind  limbs  vinaceous-tawny.  Prominent  black  spot  or  spots  along  lower 
edge  of  lateral  fold  at  its  rear  end.  Throat,  breast,  and  sides  heavily  spotted 
with  drab  or  grayish  olive  spots.  Sides  with  heavy  spots  of  hair  brown  or 
black. 

The  following  frog  came  from  Presque  Isle,  Maine,  from  Glenn  D.  Cham- 
berlain, May  15,  1946,  collected  by  Carroll  B.  Knox,  May  3,  at  the  edge  of  a 
grassy  field  adjoining  a  cedar  swamp  that  concealed  two  small  ponds  where 
wood  frogs  were  breeding. 

The  top  of  head,  upper  eyelid,  back  between  middorsal  stripe  and  dorso- 
lateral  ones  are  orange-cinnamon  or  mikado  brown  to  cinnamon-rufous  or 
tawny.  This  color  is  also  on  top  of  forearm  and  hand  and  tops  of  femur,  tibia, 
and  tarsus.  The  middorsal  stripe  is  chamois,  cinnamon-buff,  or  warm  buff, 
outlined  from  vertical  of  shoulder  backward  by  a  few  black  spots.  The  under 
edge  of  costal  fold  is  edged  with  black  and  the  sides  bear  numerous  prominent 
black  spots.  The  hind  legs  bear  three  broken  black  bars  on  tibia;  less  con- 
spicuous ones  on  femur  and  three  on  tarsus.  There  is  a  light  stripe  on  inside 
edge  of  tibia,  a  similar  one  on  outer  edge  of  tarsus,  and  one  on  femur  which 
extends  halfway  to  vent  like  the  middorsal  one.  The  mask  is  mummy  brown, 
as  are  also  the  spots  on  rear  of  brachium,  underside  of  tarsus,  stripe  from  eye 
to  snout,  and  bar  on  insertion  of  brachium.  The  stripe  on  jaw  below  mask 
extending  to  tip  of  snout  is  barium  yellow.  The  eye  is  flecked  below  with 
ochraceous-orange  and  above  with  salmon-buff.  The  underside  is  white  ex- 
cept for  underside  of  femur,  which  is  pale  ochraccous-salmon.  The  groin  and 
front  and  under  edge  of  femur  are  strontian  yellow.  There  are  slight  touches 
of  barium  yellow  on  edges  of  lower  jaw  and  on  underside  of  brachium. 

"Specimen  No.  T.  A  female  with  the  dorsal  stripe  entirely  present  and 
with  noticeable  stripes  on  the  tibia.  Throat  heavily  speckled  with  black.  When 
this  frog  was  collected,  it  was  mated  with  a  male  Rana  sylvattia.  Collected 
May  3  at  a  small  pond  ;{4  of  a  mile  north-west  of  Presque  Isle. 

"Specimen  No.  2.  A  female,  fitting  the  description  of  the  one  above  except 
for  the  speckled  throat  which  was  lacking.  This  specimen  was  also  mated  with 
a  male  Rana  syhattca  at  the  time  of  collection  and  it  laid  a  bunch  of  eggs  in  the 
school  laboratory  the  next  night.  Collected  May  $  at  the  same  pond. 

"Specimen  No.  5.  A  male  frog  with  only  about  half  of  the  dorsal  stripe  pres- 
ent. The  stripes  which  were  present  on  the  tibia  of  the  females  are  lacking 
and  there  are  only  slight  traces  of  the  speckling  on  the  throat.  Collected  May 
4  at  the  same  pond"  (letter  from  Carroll  B.  Knox,  May  13,  1947). 

The  above  is  one  of  several  justifications  for  the  practice  of  Logier  and  Toner 


548  HANDBOOK  OF  FROGS  AND  TOADS 

(forthcoming  check  list)  in  placing  all  the  wood  frogs  in  Rana  sylvatica  with- 
out subdivision  into  subspecies. 

Structure:  Body  stout;  legs  shorter  and  stouter  than  in  Rana  sylvatica;  heel 
reaching  tympanum  or  eye;  tibia  3-4  times  as  long  as  broad;  tibia  %  or  less 
than  %  the  length  of  body;  back,  flanks,  and  lower  belly  smooth;  tympanum 
smooth;  thumb  of  male  enlarged;  greater  webbing  in  hind  foot  of  males. 
The  most  apparent  difference  between  males  and  females  is  the  presence  of 
greater  webbing  in  the  hind  foot  of  males.  In  some  males  the  web  from  fifth 
to  fourth  toes  is  almost  straight,  but  in  all  it  is  convex  between  third  and 
fourth.  In  one  specimen,  the  webs  between  first  and  second  toes  are  also  con- 
vex. In  the  females,  the  edge  of  web  is  concave.  The  thumb  of  the  males  is 
enlarged,  although  not  to  the  same  extent  as  in  the  green  frog  and  meadow 
frog  groups. 

"Above  yellowish  brown.  A  dark  vitta  through  the  eye,  margined  below 
by  whitish.  Lateral  fold  of  skin  light  colored,  as  is  also  a  median  dorsal  line 
extending  from  the  snout  to  the  anus.  A  narrow  light  line  along  the  posterior 
faces  of  the  thigh  and  leg.  Tibia  half  the  length  of  body.  General  appearance 
and  size  of  R.  sylvatica.  Hab.  Cambridge,  Mass.  (Collection  of  Prof.  Agassi/.)" 
(S.  F.  Baird  [i854|  1856,  p.  62.) 

"Rana  cantabngensis  has  been  separated  from  R.  sylvatica  as  a  species  be- 
cause of  certain  color  differences  and  different  relative  measurements.  It  is 
a  smaller  and  more  stocky  frog  than  typical  R.  sylvatica.  This  is  shown  by  a 
comparison  of  the  length  of  the  tibia  with  the  total  length,  the  diameter  of 
the  tibia  with  its  length,  and  other  measurements.  Typical  cantabngensis  is 
remarkable  for  a  light  median  dorsal  stripe  and  sometimes  a  similar  light 
stripe  on  the  thigh  and  tibia.  Schmidt  and  Necker  (111.,  1933)  have  revived 
the  name  latiremis  for  the  short-legged,  stocky  frogs  from  the  north,  while, 
on  a  basis  of  the  color  characters,  they  have  referred  wood  frogs  from  Illinois, 
Indiana,  and  Michigan  to  R.  c.  cantabngcnsts  despite  the  body  proportions, 
which  are  those  of  R.  sylvatica.  The  color  characters,  as  Wright  and  Wright 
(1934)  have  already  pointed  out,  are  not  constant.  We  find  this  inconstancy 
to  be  true  not  only  for  wood  frogs  from  the  north  central  states,  but  also  for 
those  from  the  region  of  Hudson  Bay.  The  Hudson  Bay  frogs  which  lack  the 
striping  do  not  differ  structurally  from  the  striped  ones.  Color  characters  can 
not,  therefore,  be  considered  as  absolute  criteria  for  the  separation  of  the  two 
forms. 

"Since  series  of  measurements  of  frogs  from  various  localities  in  the  ranges 
of  R.  sylvatica  and  JR.  cantabngensis  demonstrate  that  there  is  no  sharp  break 
between  the  two  populations,  it  seems  evident  that  these  forms  are  best  re- 
garded as  subspecies.  For  their  differentiation,  we  prefer  the  comparative 
structural  characters  which  show  some  north  to  south  geographic  correla- 
tion, rather  than  color  characters,  which  are  apparently  variable  over  the  en- 
tire range  of  R.  cantabngensis.  Schmidt  (1938)  has  independently  come  to  a 


FROGS:  RANIDAE  549 

similar  realization  of  north  to  south  variation  in  body  form  of  these  frogs. 
We  prefer  to  give  nomenclatorial  recognition  only  to  this  variation  in  body 
form,  which  has  some  geographic  correlation,  and  not  to  color  variations 
which  apparently  lack  such  correlation. 

"R.  cantabrigensis  is  usually  separated  from  JR.  sylvatica  by  the  length  of 
the  tibia,  which  is  less  than  half  the  total  length  in  the  former  and  greater 
than  half  in  the  latter.  The  heel  extends  beyond  the  snout  in  sylvatica,  but 
does  not  reach  the  snout  in  cantabrigensis.  Specimens  from  Gaspc  Peninsula 
and  from  the  north  shore  of  the  St.  Lawrence  in  eastern  Quebec  are  somewhat 
intermediate  in  these  arbitrary  characters,  but  are  here  referred  to  S.  canta- 
brigensis" (H.  Trapido  and  R.  T,  Clausen,  Que.,  1938,  p.  123). 

"Wood-frogs  are  variable  in  color  and  markings.  For  example,  some  individ- 
uals from  western  Canada,  eastern  Canada  and  Georgetown,  Maryland  have 
spotted  breasts,  while  others  from  the  same  localities  have  no  breast  markings. 
The  upper  surface  is  also  variable,  as  mentioned  by  Wright  and  Wright  (1934) 
and  Trapido  and  Clausen  (19^8).  It  would  appear  that  the  feature  distinguish- 
ing cantabrigensis  from  sylvatica  is  length  of  hind  leg  in  comparison  to  body 
length.  The  table  shows  specimens  (Ran a  sylvatica  cantabrigensis)  from 
Yukon  to  James  Bay  and  Natashkwan  Quebec  (north  of  Anticosti  Island), 
have  the  hind  leg  shorter  than  the  body,  and  specimens  (Rana  sylvatica 
sylvatica)  with  the  hind  leg  equal  to  or  longer  than  the  body  occur  in  Nova 
Scotia,  New  Brunswick,  Gaspc  Peninsula,  southern  Quebec,  and  southern 
Ontario  westward  around  Lake  Superior.  Trapido  and  Clausen  (19^8)  refer 
specimens  from  Gaspe  Peninsula,  north  shore  of  the  St.  Lawrence  and  eastern 
Quebec  to  R.  s.  cantabngensts"  (C.  L.  Patch,  Out.,  1939,  p.  255). 

"Common  both  in  the  forest  and  at  the  water's  edge.  The  12  specimens 
collected  were  intermediate  between  typical  R.  sylvatica  sylvatica  and  typical 
R.  sylvatica  cantabrigensis  Baird,  the  heel  reaching  a  point  varying  from  the 
anterior  border  of  the  orbit  to  the  tip  of  the  snout.  None  show  the  mid-dorsal 
pale  line  commonly  found  in  cantabrigensis.  Wood-frogs  from  the  Montreal 
area  are  similar  but  have  slightly  longer  legs.  Authorities  agree  in  separating 
the  three  subspecies  of  Rana  sylvatica  (sylvatica,  cantabrigensis  t  latiremis) 
mainly  on  the  ratio  of  leg  length  to  body  length.  Data  given  by  Patch  (1959), 
Schmidt  (1958),  Boulenger  (1920),  and  others  indicate  that  when  this  cri- 
terion for  separating  sylvatica  from  cantabrigensis  is  taken  (leg  length  to 
heel  equal  to  or  greater  than  body  length  in  sylvatica)  the  territories  occupied 
by  the  two  subspecies  are  separated  by  the  St.  Lawrence  River  and  the  Great 
Lakes.  But  Preble  (1902)  and  Schmidt  (1938)  have  noted  that  there  is  a  pro- 
gressive decrease  in  the  relative  length  of  the  legs  in  the  wood  frogs  with 
increasing  latitude,  so  that  it  is  possible  that  the  differences  between  the  three 
supposed  subspecies  of  R.  sylvatica  may  be  due  to  the  influence  of  environment 
upon  the  expression  of  similar  genetic  potentialities,  R.  s.  sylvatica  of  the 
southeast  merging  into  cantabrigensis  to  the  northwest  and  the  latter  into 


550  HANDBOOK  OF  FROGS  AND  TOADS 

latiremis  in  the  far  north.  Until  further  studies  have  been  made  it  seems  ad- 
visable to  withhold  subspecific  rank  from  these  forms  of  the  wood  frog" 
(R.  Grant,  Que.,  1941,  pp.  152-153). 

"Many  seen  and  some  specimens  taken  in  Ashland  County,  one  specimen 
seen  in  Bayfield  County  about  8  miles  south  of  Ashland.  Eggs  were  found  in 
ponds  at  Camp  McLean  in  April  in  1943.  The  t/b  ratio  of  the  single  male 
taken  at  Ashland  is  .53;  in  five  females  it  varies  from  .54  to  .57,  mean  .55;  and 
in  four  juveniles  from  .50  to  .53,  mean  .51"  (R.  A.  J.  Edgren,  Wis.,  1944,  p.  497) . 

"The  proportions  suggested  by  Wright  and  Wright  (1942)  would  place  St. 
Croix  Valley  material  as  well  as  scattered  individuals  from  all  over  Minnesota 
as  belonging  to  R.  s.  sylvatica.  Patch  (1939)  has  suggested  dividing  the  species 
into  only  two  forms,  R.  sylvatica  and  R.  s.  cantabngensis  distinguishing  them 
on  the  basis  of  whether  the  leg,  minus  the  foot,  is  longer  or  shorter  than  the 
body.  On  this  basis  all  Minnesota  material  would  be  referred  to  R.  s.  canta- 
brigensis"  (W.  J.  Breckcnridge,  Minn.,  1944,  p.  91). 

We  have  always  thought  of  northern  Minnesota  wood  frogs  as  within  the 
southern  border  of  R.  s.  cantabngensis. 

Voice:  The  call  is  a  hoarse  clacking.  It  has  no  external  vocal  sac  and  calls 
from  the  surface  of  the  water. 

"In  the  early  spring  it  makes  the  ponds  resound  with  its  short,  harsh,  quack- 
ing notes"  (E.  T.  Seton,  Man.,  1918,  p.  82). 

Breeding:  This  species  breeds  from  March  to  July  depending  upon  the 
portion  of  range,  March-April  in  Michigan,  May- July  in  Mackenzie  region. 
Like  R.  sylvatica  the  eggs  are  in  masses  attached  to  vegetation.  The  tadpoles 
are  like  those  of  R.  sylvatica.  They  transform  from  May  25-Sept.  15  at  %-% 
inch  (18-22  mm.). 

E.  A.  Preble  m  "A  Biological  Investigation  of  the  Athabaska-Mackenzie 
Region"  (1908)  said  of  Ran  a  cantabngensis  latiremis  Cope:  "This  is  the  com- 
mon frog  throughout  the  region  north  to  Great  Bear  Lake  and  the  lower 
Mackenzie.  In  the  course  of  our  journeys  we  collected  a  large  series.  .  .  . 
Richardson  records  frogs,  undoubtedly  of  this  species,  from  Fort  Franklin, 
where  he  says  they  croak  loudly  in  the  beginning  of  June.  .  .  . 

"In  the  spring  of  1904  H.  W.  Jones  heard  the  notes  of  this  frog  beside 
the  Mackenzie  above  Fort  Simpson  on  April  22.  I  heard  the  first  ones  -in 
the  vicinity  of  the  post  April  28,  when  I  took  three  specimens.  At  this  time  the 
ponds  were  still  frozen  to  the  bottom  in  most  places,  but  had  thawed  in  the 
more  exposed  parts." 

Preble  has  taken  ripe  females  from  April  28  to  July  19.  There  are  records, 
however,  as  late  as  Aug.  7.  Spent  females  have  been  collected  in  May,  June, 
and  July. 

The  tadpoles  are  very  similar  to  Rana  sylvatica.  We  have  material  from 
Ontario  collected  by  Toner  and  Logier,  but  are  awaiting  tadpoles  from 
the  extreme  north  before  stating  their  characters.  The  period  is  probably  as 


FROGS:  RANIDAE  551 

variable  as  for  Rana  sylvatica,  depending  upon  early  or  late  ovulation,  and 
upon  northern  or  southern  location  in  the  range. 

There  are  records  of  transformation  from  May  25  to  Sept.  15.  The  May  and 
June  transformations  coming  in  the  same  lots  with  ripe  females  suggest  that 
possibly  some  tadpoles  winter  over,  or  some  individuals  transform  in  the  fall 
and  appear  in  the  spring  with  little  increase  in  size. 

"Rana  cantabrigensis  lattremis  Cope.  Northern  Wood  Frog.  The  most 
abundant  and  widespread  amphibian  of  the  Peace  River  district.  Spawning 
started  at  Tupper  Creek  on  May  gth.  On  this  date  the  low-pitched  croaking, 
of  remarkable  intensity  and  carrying  quality,  was  heard  at  our  camp  a  mile 
from  the  spawning  pond.  The  species  'swarms'  during  breeding  activities.  We 
found  several  aggregations  of  from  twenty  to  sixty  individuals  in  areas  of  not 
more  than  ten  square  feet  of  water  surface.  There  was  no  apparent  reason 
for  the  choice  of  these  particular  spots  on  an  homogeneous  lakeshore,  but  at 
these  points  the  egg  masses  covered  the  bottom,  lying  one  upon  the  other  in 
the  shallow  water,  quite  unattached  to  the  substratum.  After  spawning  the 
frogs  left  the  water  under  the  protection  of  the  first  rain  and  spread  out  over 
the  surrounding  country. 

"At  Swan  Lake  and  in  Tupper  Creek  these  frogs  were  very  seldom  seen 
along  the  margins  of  the  main  lake  and  the  lower  reaches  of  the  river  while 
at  Charlie  Lake  they  were  abundant  in  the  grass  along  Stodclart  Creek  and  ad- 
joining the  lake.  Possibly  the  large  numbers  of  pike  in  the  first  locality  and 
the  absence  of  this  fish  from  the  latter  water  bodies  serve  to  explain  the  dif- 
ferent behaviour  of  the  frog  population"  (I.  McT.  Cowan,  B.C.,  1959,  pp. 


In  some  of  Trapido  and  Clausen's  Quebec  material  (August,  1937)  we  have 
transformation  at  14,  16,  16,  16,  17,  17.5,  19,  20,  20  mm.  and  one  possibly  past 
at  24  mm.  In  some  of  the  Minnesota  material  (August,  1930)  appear  trans- 
formees  at  or  just  past  transformation  —  21.5,  22,  23,  25,  24,  24,  24.5,  25,  26,  26 
mm.  long.  In  some  tadpoles  from  Laurentidcs  Park,  Quebec,  we  find  tadpoles 
47,  48,  50,  50  mm.  long,  with  bodies  19.5,  20,  20,  19  mm.  The  tooth  rows  arc 
/4>  %,  %•  One  had  three  upper  rows  on  one  side  and  four  on  other  side,  and 
only  three  rows  below  with  faint  indication  of  the  fourth  row.  This  tadpole 
was  not  approaching  transformation. 

Journal  notes:  Aug.  26,  1930,  Long  Lake,  near  Hraincrd,  Minn.  Tramped 
where  grass  comes  to  sandy  shore.  The  frogs  leaped  out  onto  sand  or  piles  of 
old  sedge  stalks.  Thought  I  saw  Psetidacris  septentrionalis.  Later  concluded 
it  was  a  little  northern  wood  frog  (R.  s.  cantabrigensis).  In  the  piles  of  wet 
sedge  stalks  along  shore  found  several  small  Btifo.  By  working  grass  hard 
found  two  Hyla  cructfer,  several  small  Rana  s.  catabngensis,  and  three  Rana 
s.  cantabrigensis  half  grown.  One  of  the  half-grown  northern  wood  frogs  has 
a  beautiful  median  stripe.  Just  what  we  want.  It  also  shows  that  the  stripe  is 
not  exclusively  a  mature  coloration.  The  bulk  of  the  frogs  which  leaped  out 


552  HANDBOOK  OF  FROGS  AND  TOADS 

were  Rana  pipiens.  There  is  a  yellow  Utricularia,  Potamogeton,  sedges,  etc., 
along  the  lake  shore  edge. 

Aug.  27,  Itasca  Lake,  Minn.  Went  to  inlet  of  Itasca  Lake  in  low-lying  birch 
woods.  Thought  I'd  found  an  ideal  R.  s.  cantabrigensis  habitat — dwarf  cornel 
and  all — but  took  only  one  small  one.  About  6:30  went  with  Mr.  Olin  Coan 
to  a  spot  7  miles  away  where  a  well  had  been  dug  among  hardwood  trees.  It 
was  at  the  edge  of  a  bog  some  distance  back  from  Lake  Mallard.  There  had 
once  been  cranberries  here,  but  with  the  drilling  it  was  drying  up.  Here  next 
a  roadway  and  low  ridge,  we  caught  several  young  R.  s.  cantabrigensis  and 
one  large  one.  We  caught  about  15  small  ones,  three  being  with  median  stripes. 
One  of  the  12  without  median  stripe  had  a  light  stripe  or  area  along  either 
lateral  fold.  We  caught  one  adult — the  first  yet  taken.  The  vegetation  con- 
sisted of  many  hummocks  or  clumps  among  which  were  buck  bean,  purple 
cinquefoil,  wild  calla,  grass  of  parnassus — really  a  bog  situation.  Around  this 
area  were  many  alders  (Alnus). 

Aug.  28.  Toward  Bemidji  went  through  a  tamarack  woods.  Ditches  either 
side  of  the  road,  took  several  adult  R.  s.  cantabrigensis  and  young.  In  tamarack 
woods  noticed  bunchberry,  Dahbardia  repens,  mountain  maple,  buckthorn, 
Chiogenes,  and  other  bog  forms.  Later  along  road  found  spot  where  road  had 
sunk  into  the  swamp.  Here  took  a  good  male  northern  wood  frog.  It  is  surely 
the  form  of  these  ditches  along  these  highways  of  the  tamarack  region.  Only 
one  or  two  had  the  median  dorsal  stripe. 

Aug.  30,  Pembina,  N.D.,  Red  River  of  the  North.  Late  in  the  afternoon 
went  down  to  the  river.  The  river  is  low  with  a  zone  of  6  feet  of  mud  inclined 
and  exposed.  This  is  broken  into  blocks  %-iK»  feet  and  is  clear  of  vegeta- 
tion. Cracks  1-5  inches  wide.  Blocks  6-9  inches  deep.  Then  comes  a  zone  of 
low  weeds,  dandelions,  some  small  amaranthlike  plant,  and  very  low  grasses. 
Then  area  of  dense  low  willows  up  bank  for  2  rods  or  more,  very  dry  and 
baked.  Then  upland  meadow  or  cultivated  field.  Here  by  walking  along  moist 
edge  of  vegetation  wood  frogs  and  meadow  frogs  leaped  out  onto  the  mud 
blocks.  Took  several  median-striped  northern  wood  frogs.  For  the  first  time 
found  the  median  stripe  predominant.  Took  adult  male  with  median  stripe 
halfway  to  vent,  a  male  without  median  stripe,  a  female  with  median  stripe 
from  snout  to  vent,  and  a  female  without  median  stripe.  Several  young  with 
and  several  without  median  stripe.  All  quite  spotted  on  breast.  Dark  pattern 
on  back  quite  conspicuous.  This  is  the  finest  series  of  northern  wood  frogs 
we  have  taken  or  seen  alive. 

Authorities'  corner: 

G.  A.  Boulenger,  Gen.,  1891,  p.  453.  G.  A.  Boulenger,  Gen.,  1920,  pp.  457- 

R.  H.  Howe,  Jr.,  Gen.,  1899,  p.  374.  458. 

E.  A.  Treble,  Kcewatin,  1902,  pp.  133-     C.  L.  Patch,  B.C.,  1922,  p.  77. 

134. 


FROGS:  RANIDAE  553 

Mexican  Frog 

Rana  tarahumarae  Boulenger.  Plate  CXXII;  Map  35. 

Range:  "Sierra  Tarahumarae,  N.  W.  Mexico  about  3000  feet"  (Boulenger, 
Gen,,  1919,  p.  415);  Pefia  Blanca  Spring,  Santa  Cruz  Co.,  Ariz.;  i  mile  above 
the  XSX  Ranch,  east  fork  of  the  Gila  River,  Socorro  Co.,  N.  Mex.  (J.  M. 
Linsdale,  N.  Mex.,  1933,  P-  222)- 

"Mcsilla  Valley,  R  M."  (Little  and  Keller,  N.  Mex.,  1937,  P-  221).  "Rose 
Creek,  Roosevelt  Reservoir  Area,  Ariz.  1940"  (E.  L.  Little,  Jr.,  Ariz.,  1940, 
p.  262) . 

Habitat:  "The  colony  was  clustered  around  a  series  of  pot-holes  in  a  can- 
yon, ,  .  .  and  no  running  water  could  be  found  in  the  region.  Besides  the 
twenty  or  so  adults  seen  in  these  pot-holes,  a  group  of  probably  half  that 
number  was  found  in  an  old  tumbled-m  mine  which  had  filled  with  water" 
(B.  Campbell,  Ariz.,  1931,  p.  164). 

"All  [were]  in  small  pools  bordered  by  willows  and  cotton  woods  along  Rio 
Grande  near  Mesilla  Dam"  (E.  L.  Little,  Jr.,  and  J.  G.  Keller,  N.  Mex.,  1937, 
p.  221). 

"A  series  of  22  specimens  was  taken  out  of  about  100  observed  in  pools  of 
water  shaded  by  alders  and  sycamores  along  Rose  Creek,  elevation  about  5400 
feet,  at  the  low  border  of  the  pine-fir  zone,  on  August  22,  1936"  (E.  L.  Little, 
Jr.,  Ariz.,  1940,  p.  262) . 

Size:  Adults,  zYs-qYz  inches  (58-113  mm.).  "The  resemblance  of  the  adults 
to  Rana  boylii  is  very  noticeable,  but  the  size  is  distinctive — the  three  speci- 
mens on  hand  measuring  81, 91  and  1 13  mm.  (head  and  body)"  (B.  Campbell, 
Ariz.,  1931,  p.  164). 

"One  specimen  70  mm.  long  was  caught  on  August  n,  1934  and  9  smaller 
ones  averaging  50  mm.  in  length  were  taken  November  4,  1934"  (E.  L. 
Little,  Jr.,  and  J.  G.  Keller,  N.  Mex,,  1937,  P-  221). 

General  appearance:  This  is  a  large  dusky-olive  frog,  with  a  few  black  spots 
on  the  back  and  broken  crossbands  and  dark  spots  on  the  legs,  chunky  of 
body  and  broad  in  the  head  with  rounded  snout. 

Color:  Female.  Pefia  Blanca  Spring,  Ariz.,  June  19,  1934.  The  upper  parts 
are  lincoln  green  or  dusky  olive-green.  There  are  four  black  or  dusky  olive- 
green  transverse  bars  on  femur  to  the  knee,  four,  sometimes  broken,  bands 
on  tibia  and  two  on  tarsus,  and  several  on  outer  edges  of  foot  and  toes;  these 
all  distinct.  Back  is  marked  with  many  small,  round,  black  or  olivaceous  black 
spots.  These  also  show  on  top  of  head  to  level  of  eyes.  Top  of  head  and  face 
is  buflEy  olive  with  numerous  round  spots  of  isabella  color.  The  tympanum  has 
color  of  back.  On  one  side  is  quite  a  prominent  olive-ocher  or  ecru-olive  band 
from  in  front  of  eye  to  angle  of  mouth,  becoming  back  of  V  of  mouth  honey 
yellow.  On  other  side  it  is  almost  absent,  being  olive-ocher  just  in  front  and 
underneath  the  ear.  Side  of  body  is  largely  olive-ocher.  Under  parts  white, 


554  HANDBOOK  OF  FROGS  AND  TOADS 

except  for  throat  which  is  heavily  clouded.  Rear  of  femur  has  some  clay  color 
among  the  dark  spots.  Underside  of  tibia  and  front  of  foot  with  some  honey 
yellow.  Iris  ring  citron  yellow.  Iris  olivaceous  black  with  spottings  of  light 
russet-vinaceous. 

Structure:  The  male  has  a  swollen  thumb  with  a  slight  tendency  toward 
a  diagonal  oblique  depression  across  its  middle  like  1?.  boylii  subspecies. 

"Vomerinc  teeth  in  small  groups  close  together  behind  the  level  of  the 
choanae. 

"Head  broader  than  long,  much  depressed;  snout  rounded,  feebly  project- 
ing beyond  the  mouth,  as  long  as  the  eye;  canthus  rostralis  indistinct;  loreal 
region  very  oblique,  slightly  concave;  nostril  equidistant  from  the  eye  and 
from  the  tip  of  the  snout;  distance  between  the  nostrils  equal  to  the  inter- 
orbital  width,  which  is  equal  to  or  a  little  less  than  that  of  the  upper  eyelid; 
tympanum  distinct,  without  or  with  a  few  small  asperities,  %  to  %  the 
diameter  of  the  eye,  once  to  once  and  a  half  its  distance  from  the  latter. 

"Fingers  moderate,  the  tips  feebly  swollen,  first  longer  than  the  second, 
third  longer  than  the  snout;  subarticular  tubercles  large,  prominent. 

"Hind  limb  long,  the  tibio-tarsal  articulation  reaching  the  tip  of  the  snout, 
the  heels  meeting  when  the  limbs  are  folded  at  right  angles  to  the  body;  tibia 
4  to  4%  times  as  long  as  broad,  i%  to  2  times  in  length  from  snout  to  vent, 
shorter  than  the  fore  limb,  as  long  as  or  slightly  longer  or  shorter  than  the 
foot.  Toes  with  the  tips  swollen  into  small  disks,  the  base  of  which  is  in- 
volved in  the  very  broad  web;  outer  mctatarsals  separated  nearly  to  the  base; 
subarticular  tubercles  rather  large  and  prominent;  no  tarsal  fold;  inner  meta- 
tarsal  tubercle  elliptic,  feebly  prominent,  %  to  %  the  length  of  the  inner  toe; 
no  outer  tubercle. 

"Skin  smooth,  or  upper  parts  with  small  pustules;  a  feeble,  curved  glandular 
fold  from  the  eye  to  the  shoulder"  (G.  A.  Boulengcr,  1920,  p.  468) . 

Breeding:  "The  bleeding  season  is  evidently  after  the  heavy  summer  rains 
which  begin  in  July"  (B.  Campbell,  Ariz.,  1931,  p.  164). 

The  tadpole  is  quite  large,  ?%~4  inches  (88-101  mm.),  and  the  tooth  ridges 
are  %,  %.  On  June  21,  1954,  Wlt^  Mr.  G.  W.  Harvey  we  took  many  mature 
tadpoles  at  Alamo  Canyon,  Pena  Blanca,  Ariz.,  and  some  at  transformation. 
The  tadpoles  averaged  between  70-88  mm.  and  transformed  at  30-35  mm. 
B.  Campbell  found  them  transforming  June  18,  1931. 

In  general  appearance  the  tadpole  is  deep  green  with  very  close-set  small 
spots  on  the  body.  These  become  much  larger  and  prominent  on  tail  muscula- 
ture and  tail  crests.  The  body  is  buffy  citrine  or  yellowish  olive  to  dark  green- 
ish olive.  The  spots  may  be  black,  brownish  olive,  or  olive.  In  some  tadpoles 
the  ground  color  is  so  dark  that  no  dorsal  spotting  is  revealed.  In  lighter- 
colored  tadpoles  the  small  spots  are  quite  prominent  on  sides  and  top  of  body 
and  top  of  head;  the  face  usually  has  less  spotting.  The  belly  and  throat  are 
white  with  the  lateral  spots  encroaching  on  either  side.  That  color  may  be 


FROGS:  RANIDAE 


555 


like  the  background  color  or  bone  brown.  There  is  a  wash  along  the  sides 
of  olive  lake,  as  one  gets  toward  the  developing  hind  legs.  Occasionally  in  the 
tadpole  there  is  a  reminder  on  rear  upper  edge  of  the  labial  of  an  olive-ocher 
line  suggestive  of  the  yellow  line  of  the  jaw  of  the  adult.  The  tail  has  much 
of  same  background  color  as  the  back,  being  conspicuously  spotted  with  large 
black  or  brownish  olive  spots.  The  upper  and  lower  tail  crests  are  as  con- 
spicuously spotted  as  the  tail  musculature,  except  the  basal  third  or  fourth 
of  lower  tail  crest,  which  is  free  of  spots.  Toward  tip  of  tail  strongly  washed 
with  Hay's  russet,  apricot  buff,  or  ochraceous  buff  in  interspaces.  The  ventral 
side  of  musculature  on  either  side  of  clear  portion  of  under  tailcrest  is  with- 
out spots,  but  the  paired  veins  on  either  side  stand  out  prominently.  The  iris 
is  dark  olive  with  citrine  yellow  rim,  dotted  with  apricot  buff  and  olive  lake. 
Teeth  %.  Hind  legs,  on  upper  portions,  one  shade  lighter  than  ground  color. 

When  we  first  glanced  at  the  tadpoles  they  did  not  impress  us  as  of  the 
R.  boylii  group,  but  somehow  (possibly  irrelevantly)  like  the  R.  grylio-R. 
virgatipes-R.  hec1(scheri  group.  Of  couise  they  are  not  marked  like  R.  hec^- 
schen  but  the  tadpoles  are  marked  as  conspicuously  in  a  different  way. 

Journal  notes:  June  19,  1934,  Pefia  Blanca  Spring,  Ariz.  Last  night  saw  a 
Rana  tarahttmarae  on  weight  stone  of  the  spring's  concrete  tank.  Saw  another 
near  the  outlet.  Caught  one  near  overflow  pipe.  Yes,  it  is  Berry  Campbell's 
/?.  tarahttmarae. 

June  21,  TQ34-  Went  at  6  A.M.  with  Mr.  G.  W.  Harvey  through  Aqua  Fria 
Canyon  to  Alamo  Canyon  where  he  first  showed  Campbell  the  frogs.  It  is  a 
narrow  canyon.  Did  not  try  tumbled-in  mine.  In  first  pool  saw  a  large  one 
and  no  end  of  striking  spotted  tadpoles.  Took  several  with  dip  net;  we  could 
bring  it  in  water  right  up  to  the  frog,  and  he  would  jump  into  it.  Could  see 
large  ones  on  bottom  and  bring  net  up  to  them.  One  large  male  in  the  water 
looked  to  have  bronzy  or  brick  red  casts  on  the  legs  and  elsewhere.  Their 
webbed  fourth  toe  indicates  they  are  quite  aquatic.  In  shallower  pools  R. 
piptens  adults  and  tadpoles.  Most  of  the  bigger  pools  had  only  R.  tarahtimarae. 
Took  a  series  from  tadpole  to  transformation  and  secured  ten  adults.  Could 
have  taken  many.  They  certainly  are  Rana  £oy///-hke.  Are  they  nearest  to 
R.  b.  sierrae  or  R.  b.  rntiscosa?  The  males  have  clouded  throats.  When  Camp- 
bell was  here  the  water  was  much  higher. 

Campbell's  largest  adult  with  its  uniform  color,  cloudy  throat  and  venter, 
fully  webbed  hind  feet  (almost  straight  from  tip  of  fifth  toe  to  fourth  toe) 
looked  as  much  a  member  of  the  Rana  hec{scherif  R.  catesbeiana,  R.  virgatipes 
type  as  of  the  R.  boyhi  group.  Examination  of  the  smaller  examples  bring 
out  better  the  R.  boylii  resemblances. 

Authorities'  corner: 

B.  Campbell,  Ariz.,  1931,  p.  164.  E.  L.  Little,  Jr.,  and  J.  C.  Keller,  N. 

J.    M.    Linsdalc,    N.    Mex.,    1933,    p.         Mex.,  1937,  p.  221. 

222.  E.  L.  Little,  Jr.,  Ariz.,  1940,  p.  262. 


556  HANDBOOK  OF  FROGS  AND  TOADS 

Sphagnum  Frog,  Carpenter  Frog,  Cope's  Frog 

Rana  virgatipes  Cope.  Plate  CXXIII;  Map  31. 

Range:  New  Jersey  to  Okefinokee  Swamp,  Ga.  Delaware  (Conant).  South 
Carolina  (Chamberlain,  1939). 

Habitat:  This  species  may  be  in  the  sphagnum  edge  of  open  ponds,  in 
sphagnum  mats  in  deeper  parts  of  ponds  or  lakes,  in  wooded  edges  of  some 
of  the  coastal  rivers,  in  branch  swamps  of  the  Southeast,  in  wooded  inlets 
or  outlets  of  open  ponds,  cypress  bays,  or  strands  of  islands  in  southern 
swamps,  or  at  times  in  the  open  water-lily  prairie. 

Size:  Adults,  i%-2%  mches.  Males,  41-63  mm.  Females,  41-66  mm. 

General  appearance:  This  small  frog  has  a  long,  narrow  head,  back  brown- 
ish with  four  yellowish  or  golden-brown,  longitudinal  stripes.  The  under  parts 
are  yellowish  white  with  dark  brown  or  black  spots.  The  rear  of  the  femur 
has  alternating  dark  and  light  stripes.  The  sides  also  are  marked  with  black- 
ish spots. 

In  coloration  JR.  virgatipes  looks  most  like  R.  gryho.  In  adult  size  it  is  more 
like  R.  septentrionahs  but  smaller,  or  is  possibly  like  a  small  R.  catesbeiana. 
At  Lakehurst  Lake  it  reminds  one  of  the  water  prairie  form,  R.  grylio;  in  the 
sphagnum  strands  and  thickets  of  the  Okefinokee  it  reminds  me  of  the  peat- 
lake  species  of  Canada  or  beaver-lake  thicket  form  of  the  Adirondacks,  namely, 
R.  septentrionalis.  In  egg  mass  it  is  most  like  R.  septentrionalis,  but  in  individ- 
ual eggs  unlike  R.  septentrionalis \  R.  clamitans,  and  R .  grylio  because  it  has  no 
inner  jelly  envelope.  The  individual  eggs  are  more  like  the  eggs  of  the  film 
species  R.  catesbetana,  but  the  jelly  is  firmer.  The  tadpole  reminds  one  most  of 
R"  gryho  in  coloration,  but  it  is  more  of  the  R.  clamitans  type.  Like  all  of  the 
R.  clamitans  and  R.  catesbetana  type  it  winters  over  as  a  tadpole.  It  transforms 
at  a  size  near  that  of  R.  clamitans  and  may  have  a  larger  tadpole  than  that 
species.  Therein  it  approaches  1?.  catesbeiana,  R.  grylio,  and  R.  septentrionalis. 

Color:  Male.  Okefinokee  Swamp,  Ga.,  July  15,  1921.  Back  natal  brown  or 
chestnut  brown  with  black  spots.  Natal  brown  on  dorsum  of  fore  and  hind 
legs.  Stripe  down  either  side  from  eye  to  hind  leg  hazel  or  fawn  color.  Smaller 
stripe  of  light  or  pale  ochraceous-salmon  from  angle  of  mouth  halfway  down 
side.  This  stripe  continues  over  angle  of  mouth  and  on  to  upper  jaw  as  pale 
greenish  yellow.  Throat  sulphur  yellow  with  indefinite  deep  brown  spots. 
Rest  of  under  parts  cartridge  buff  to  pale  chalcedony  yellow  with  beautifully 
marked  bone  brown  or  almost  black  spots  on  venter  and  sides.  Back  and  for- 
ward parts  of  iris  neva  green,  rim  green  yellow.  Iris  proper  spotted  black  and 
empire  yellow. 

Female.  Okefinokee  Swamp,  Ga.,  July  22,  1922.  Back  olive-brown.  Back 
stripe  and  side  stripe  buffy  brown.  Throat  sulphur  yellow,  also  inner  edge  of 
fore  limb.  This  yellow  also  from  angle  of  mouth  to  over  arm  insertion.  Spot- 
ting on  under  parts  not  so  prominent  as  in  male  nor  so  clear-cut,  but  it  is 


FROGS:  RANIDAE 


557 


Plate  CXX1L  Kana  tarahumarae 
lti&7*  Males.  2.  Tadpole.  4.  Female.  6. 
Transformed  frog. 


Plate  CXXUL  Rana  virgatipcs.  1,3.  Fe- 
males (X4/o).  2,5.  Males  (X%).  4.  Male 
croaking  (X%). 


558  HANDBOOK  OF  FROGS  AND  TOADS 

conspicuous.  Females  in  general  look  white  on  under  parts,  while  males  have 
washes  of  yellow  on  throat,  on  sides,  and  slightly  on  other  parts.  Eye  as  in  the 
male. 

Structure:  Tympana  enlarged  in  the  males,  but  not  so  striking  as  in  Rana 
grylio,  R.  catesbeiana,  R.  damitans,  or  R.  fisheri  (except  in  very  old  males), 
nor  is  the  thumb  quite  so  large  proportionally;  no  dermal  folds;  two  joints  of 
fourth  toe  free;  males  with  vocal  pouch  on  either  side;  these  being  perfectly 
round  vesicles  when  inflated,  and  the  throat  not  inflating. 

"Rana  virgatipes,  sp.  nov. — Vomerme  tooth  patches  between  the  choanae, 
and  extending  posteriorly  to  their  posterior  border.  Hind  legs  short,  the  heel 
extending  from  the  middle  of  the  tympanum  in  some  to  near  the  eye  in  other 
specimens.  Webs  rather  short,  two  phalanges  of  the  fourth  toe  free.  Praehallux 
small,  but  quite  prominent;  no  external  solar  tubercle.  Skin  of  upper  surfaces 
of  body  and  posterior  limb  covered  with  minute  tubercles;  no  longitudinal 
dermal  folds.  Males  with,  females  without,  external  vocal  vesicles.  Interocular 
width  one-half  that  of  each  eyelid.  Tympanic  disk  distinct,  equaling  the  eye 
in  longest  diameter.  Head  (to  posterior  border  of  tympanic  disk)  about  one- 
third  length  of  head  and  body.  End  of  muzzle  oval-acuminate,  projecting 
moderately  beyond  mouth  border.  Nostril  opening  vertically  equidistant  be- 
tween border  of  orbit  and  end  of  muzzle.  First  and  second  fingers  subequal 
and  longer  than  the  fourth. 

"Length  of  head  and  body,  60  mm.;  width  of  head  at  posterior  borders  of 
tympana,  21.5  mm.;  length  of  fore  limb  from  axilla,  28  mm.;  length  of  hind 
limb  from  vent,  76  mm.;  length  of  hind  foot,  39  mm.;  of  tarsus,  15  mm.;  of 
tibia,  23  mm."  (E.  D.  Cope,  N.J.,  1891,  pp.  1017-1018). 

Voice:  The  call  is  like  the  blow  ot  a  hammer,  usually  repeated  three  to  six 
times  in  rapid  succession,  or  like  wood  choppers. 

W.  T.  Davis  (1907)  said,  "Its  note  sounds  much  like  the  blow  of  a  hammer 
on  a  board.  It  is  a  quickly  uttered  Chuc\-up,  chucl^up  and  the  frog  usually 
hammers  three  or  four  times."  Fowler  likens  it  to  "the  noise  produced  by 
wood  choppers  cutting  trees  a  short  distance  back  in  the  forest."  Harper  calls 
it  "a  rather  loud,  incisive,  distinctive,  clucking-croak."  The  croak  is  four  to 
seven  notes  in  rapid  succession,  all  alike.  Each  part  occupies  a  second  or  less 
in  duration.  Besides  this,  the  species  has  another  call  of  two  notes  lower  Jn 
pitch.  The  chorus  starts  about  dusk  or  sometimes  before  and  gathers  strength 
as  midnight  approaches.  At  3  or  4  in  the  morning  they  go  strong.  At  daybreak 
one  to  several  may  be  heard  and  these  may  continue  until  8:30  to  9:00  A.M. 
Once  in  a  while  it  is  heard  by  day  during  cloudy  weather. 

"The  clack,  clack,  clack,  clack,  clack  of  this  frog  could  be  heard  about  the 
Rancocas  Creek  above  New  Lisbon,  in  Burlington  county,  on  May  i2th,  1907. 
It  occurred  at  intervals,  interrupting  the  stillness  of  a  backward  spring. 
Though  loud  and  not  often  uttered,  in  comparison  with  those  found  at  Mare 
Run,  their  croaking  could  be  heard  at  quite  a  distance.  Sometimes  the  animals 


FROGS:  RANIDAE 


559 


must  have  been  quite  close,  for  we  could  hear  them  when  but  a  few  feet 
away  without  seeing  them.  It  may  have  been  that  the  weather  was  too  cold, 
for  their  croaking  was  always  located  as  coming  from  among  the  submerged 
and  overgrown  vegetation  along  the  banks.  In  such  places  the  temperature 
was  considerably  higher  by  midday  than  elsewhere.  When  Mr,  Hunt  visited 
this  place  just  a  year  previously,  the  frogs  were  very  numerous.  The  weather 
at  that  time,  however,  though  about  the  same  time  in  May,  was  much  warmer. 
In  the  evening,  during  the  night,  and  in  the  early  morning,  they  were  very 
noisy,  but  during  the  day  were  more  or  less  quiet,  only  an  occasional  croak 
being  heard  at  intervals,  or  as  noted  above"  (H.  W.  Fowler,  N.J.,  1908,  p.  194). 

Breeding:  This  species  breeds  from  late  April  to  mid-August.  The  egg  mass 
is  a  plinth  3-4  inches  (75-100  mm.)  by  i%  inches  (38  mm.),  or  it  may  be 
globular.  The  complement  is  small,  200-600  eggs.  The  black  and  creamy-white 
eggs  are  large  and  far  apait.  The  egg  is  Mo  (1.5-1.8  mm.),  the  envelope 
%-l/\  inch  (3.8-6.9  mm.).  The  tadpole  is  large,  3%  inches  (92  mm.),  dark 
in  color  with  a  few  black  spots,  its  tail  grayish  with  a  row  of  large  spots  in 
the  upper  crest.  The  tooth  ridges  arc  %  or  Mi-  After  a  tadpole  period  of  i  year, 
the  tadpoles  transform  in  early  spring  at  ll%<i-i!4  inches  (23-31  mm.). 

This  frog  passes  the  winter  as  a  tadpole  and  transforms  in  early  spring  be- 
fore May  15,  a  few  stragglers  going  along  to  mid-July.  C.  S.  Brimley  in  North 
Carolina,  W.  T.  Davis  and  we  at  Lakehurst,  N.J.,  in  May,  and  T.  and  F.  K. 
Barbour  at  Lakehurst  in  July  have  taken  specimens  33-37  mm.  just  beyond 
transformation. 

Journal  notes:  On  May  22-23,  1924,  we  heard  a  few  males  croaking  on  the 
south  side  of  the  west  point  into  the  lake  at  Lakehurst,  N.J.  It  was  noon  and 
the  sun  was  shining.  Here  in  the  sphagnum-heath  edge,  we  caught  four 
adults,  two  females  and  two  males.  As  we  waded  along  we  would  see  them 
sometimes  wholly  out  of  water  or  at  the  lake's  edge.  Usually  they  leaped  into 
the  water  and  hid  under  the  vegetation  mat  or  quickly  came  up  under  a  water- 
lily  leaf  or  swam  some  distance  and  then  poked  out  their  heads.  Most  of  the 
males  were  out  in  the  deeper  water. 

On  Black  Jack  Island,  July  28,  1921,  we  had  one  of  the  hardest  rams  I  ever 
experienced.  In  the  morning,  July  29,  with  a  temperature  of  73°,  I  heard  a 
carpenter  frog  chorus.  They  were  scattered  in  among  the  small  cypress  where 
there  was  an  undercarpet  of  Woodwardta  and  a  little  sphagnum.  They  seemed 
to  be  around  the  bases  of  the  cypress  trees  and  on  the  cypress  knees.  I  saw 
none  of  them.  In  the  daytime  they  are  shy. 

In  1922  we  camped  the  night  of  June  9  on  the  north  side  of  Everett's  Pond 
(about  5  or  10  min.  ride  by  auto  from  the  N.C.-S.C.  state  line),  N.C.  In  the 
wooded  sections  where  streams  flowed  into  the  lake  in  dense  thickets  of  Smilax 
(briar)  and  other  plants  were  several  Rana  virgatipes  croaking.  In  1922  on 
June  21  in  the  prairie  just  north  of  Lake  Sego  in  the  Okefinokee  Swamp,  heard 
seven  or  eight  R.  virgattpes.  Found  one  in  a  grassy  place  with  water  lilies. 


560  HANDBOOK  OF  FROGS  AND  TOADS 

On  a  water-lily  pad  was  the  one  I  captured.  They  like  grassy  places  near  the 
edge  of  a  head  (islet)  which  is  bordered  with  sphagneous  strand.  Also  seem 
to  like  some  timbered  parts. 

June  8, 1929.  Needed  a  male  and  female  Rana  virgatipes  for  photographs. 
Boarded  a  train  at  Ridgewood  for  Lakehurst  and  without  collecting  kit  or 
proper  clothes.  It  rained  until  midafternoon.  With  every  handicap,  we  waded 
the  edge  of  the  lake  with  no  success.  The  same  result  along  the  west  point  of 
landing  extending  into  the  lake.  The  vegetation  mats  have  not  reached  the 
surface  to  any  appreciable  degree.  Mr.  Shinn  and  his  boys  have  a  cottage 
at  the  base  of  the  point.  They  informed  us  that  locally  the  frogs  have  long 
been  called  "carpenter  frogs." 

Some  new  visitors  beside  the  lake  can  hardly  sleep  at  night  when  the  species 
is  in  full  chorus.  Mr.  Shinn  sent  the  boys  out  in  a  canoe  to  get  some  on  the 
mats  in  mid-lake  and  along  the  east  side.  We  went  to  Emslie's  Pond.  When 
we  returned  the  boys  had  four  males,  one  female,  and  one  recently  trans- 
formed frog.  Several  they  had  thrown  back.  They  approached  lily  pads  and 
with  a  scoop  of  a  long-handled  dip  net  caught  the  frogs.  The  lily  pad  and 
mid-lake  habits  suggest  Rana  septentrionalis  of  the  Adirondack  Mountains 
and  the  North. 

Authorities'  corner: 

W.  T.  Davis,  N.J.,  1907,  pp.  49,  50.        C.  S.  Brimley,  N.C.,  19273,  p.  12. 
H.  W.  Fowler,  N.J.,  1908,  pp.  191-     E,  B.  Chamberlain,  S.C.,  1939,  p.  34. 

195.  R.  Conant,  Del.,  1945,  p.  4. 


FAMILY  BREVICIPITIDAE 

Genus  HYPOPACHUS  Keferstein 
Map  37 

The  best  recent  discussion  of  the  genus,  that  by  E.  H.  Taylor  (Tex.,  19400, 
pp.  515-516),  makes  any  other  evaluation  useless.  Following  his  remarks  on 
the  genus,  Taylor  describes  five  new  forms,  prefacing  them  with  a  good  ap- 
praisal of  Hypopachus  cuneus  cuneus  (pp.  517-528).  Before  Taylor  three  more 
had  been  recogni/ed,  making  at  least  13  now  known  in  the  genus. 

Taylor's  Toad 

Hypopachus  cuneus  Cope.  Plate  CXXIV;  Map  37. 

Range:  Southern  Texas  southward,  Tamaulipas,  Mexico  (Smith-Dunkle) . 

Habitat:  Subterranean  form.  We  do  not  know  whether  or  not  it  breeds  in 
such  transient  pools  as  Microhyla.  Our  own  field  experience  is  restricted  to 
tadpoles  and  transforming  frogs. 

"Adults  driven  from  under  mesquite  trees  during  irrigation  were  found 
to  have  been  feeding  on  termites  and  minute  dipterous  insects.  Specimens  in 
captivity  likewise  fed  readily  upon  termites.  Even  though  buried  several 
inches  underground,  while  in  captivity,  the  placing  of  termites  upon  the  sur- 
face readily  brought  them  into  the  open.  In  the  natural  state,  however,  they 
do  not  come  into  the  open  except  when  driven  out  by  excessive  rains.  They 
invariably  are  to  be  found  in  burrows,  among  the  trash  of  pack  rats,  and  in 
the  hollows  under  trees.  As  humidity  decreases,  they  apparently  seek  out 
deeper  and  more  moist  situations,  or  burrow  backwards  into  the  soil,  where 
they  are  not  subjected  to  rapid  drying  out"  (S.  Mulaik  and  D.  Sollberger,  Tex., 
1938,  p.  90). 

Size:  Adults,  1-1%  inches.  Males,  smaller,  25.0-37.5  mm.  Females,  29- 
41  mm. 

General  appearance:  The  conspicuous  mark  is  the  light  yellow  or  orange 
thread  stripe  down  mid-back.  A  similar  white  line  extends  length  of  gray- 
mottled  venter  with  a  branch  to  each  arm.  Broad  constant  oblique  white  mark 
from  eye  to  angle  of  mouth,  thence  to  the  arm  insertion.  This  line  bordered 
by  black  spots.  An  irregular  line  of  black  dots  may  extend  from  middorsal 
line  near  head  to  upper  groin  on  either  side.  The  sides,  groin,  rear  of  femur, 


I 


NARROW-MOUTH  TOADS:  BREVICIPITIDAE 


563 


and  sides  of  legs  are  heavily  marked 
with  inky  spots.  There  is  a  cinna- 
mon, fawn,  or  orange  tinge  in  the 
groin,  on  the  femur,  and  on  the  up- 
per arm. 

Color:  One  female  and  five  males, 
Kingsville,  Texas,  from  C.  L.  Reed, 
August,  1931. 

Female.  Upper  parts  of  back  brus- 
sels  brown  or  snuff  brown  with  same 
color  on  top  of  head,  upper  surface 
of  tibia,  rear  edge  of  tarsus,  and  hind 
foot  to  base  of  fifth  toe.  Beginning 
%  inch  from  transverse  head  fold, 
there  goes  off  from  dorsal  line  a  row 
of  black  dots,  which  extend  irregu- 
larly to  the  upper  groin.  The  light 
line  down  middle  of  back  is  orange- 
buff.  Just  above  vent  is  a  very  nar- 
row transverse  orange-buff  line  lead- 
ing to  rear  angle  of  knee,  then 
broken,  to  be  resumed  later  halfway 
down  on  rear  of  tibia  and  on  to  the 
tarsus,  where  it  becomes  whitish. 
From  the  point  where  these  lines 
meet  the  dorsal  line,  the  dorsal  ex- 
tends caudad  to  vent  as  a  white  line. 
The  tip  of  snout  and  canthus  are 
olive-buff  or  deep  olive-buff,  with  a 
black  border  from  the  eye  to  tip  of 
snout  below  this  color.  This  snout 
color  breaks  the  continuous  dorsal 
line.  On  the  ventral  side  of  snout, 
this  median  line  resumes  as  a  short 
white  line.  Broken  by  the  mouth,  it 
continues  as  a  white  line  down  mid- 
dle of  venter  to  end  of  belly.  On  the 
pectoral  region  a  diagonal  white  line 
goes  to  the  axilla  and  continues  on 
the  rear  of  fore  limb  to  the  base  of  the  hand.  The  dorsal  color  is  bordered  back 
of  eye  to  groin  and  on  either  side  of  hind  legs  by  a  more  or  less  broken  string 
of  irregular  black  blotches,  which  are  particularly  prominent  in  the  groin  and 
on  rear  of  femur.  The  black  spots  of  groin,  hind  legs,  and  forelegs  are 


Plate  CXX1V.  Hypopachus  cuneus.  1,3. 
Female  (X*%)-  2.  Male  (X^)- 


564  HANDBOOK  OF  FROGS  AND  TOADS 

interspersed  with  rufous,  flesh  ocher,  onion-skin  pink,  or  light  vinaceous- 
cinnamon  tinges.  The  stripe  from  eye  obliquely  down  to  an  area  intermediate 
between  forearm  insertion  and  mouth  is  white,  irregularly  bordered  on  either 
side  with  black  spots.  That  portion  between  eye  and  level  of  mouth  is  tinged 
with  cinnamon-buff.  The  sides  from  white  oblique  stripe  to  groin  are  almost 
solid  smoke  gray  or  light  grayish  olive.  On  the  belly  this  becomes  palp  smoke 
gray  mottled  with  drab-gray,  while  the  throat  becomes  more  light  drab  than 
smoke  gray.  Iris  of  one  side  brazil  red  or  flame  scarlet  heavily  speckled  with 
black.  Iris  rim  is  pale  green-yellow.  Iris  of  other  side  green-yellow  heavily 
spotted  with  black  with  iris  rim  sulphur  yellow.  Pupil  round. 

Male.  The  dorsal  color  is  Saccardo's  olive  to  citrine-drab  with  an  oblique 
line  of  obscure  black  dots  from  median  line  at  shoulder  to  groin.  The  groin, 
rear  and  front  of  femur,  and  sides  are  more  heavily  spotted  with  black  than 
in  female,  although  the  lateral  black  band  is  more  prominent  in  the  female. 
The  white  mid-ventral,  pectoral,  brachial,  and  antebrachial  stripes  are  less 
prominent  than  in  female.  The  stripe  down  back  is  very  prominent,  but  the 
dorsofemoral  lines  are  less  prominent.  The  light  dorsal  lines  are  buff-yellow 
or  light  orange-yellow.  The  bright  color  of  groin  and  concealed  surfaces  of 
hind  legs  is  not  so  bright  in  the  five  males  in  hand  as  in  the  one  female  that 
we  have.  This  bright  tinge  is  cinnamon  to  tawny-olive.  The  color  of  top  of 
forearms  and  hind  limbs  of  two  is  isabella  color.  In  one,  the  light  diagonal 
band  is  wholly  cream-buff;  in  another,  upper  end  cream-buff  and  lower  end 
cartridge  buff.  The  mottling  of  the  venter  is  much  heavier  and  more  strongly 
contrasted  than  in  female.  The  throat  is  blackish  slate,  plumbeous-black,  or 
dark  quaker  drab.  From  end  of  light  oblique  stripe  across  to  other  end  is  the 
rear  margin  of  this  dark  area.  Behind  this  dark  area  is  an  area  of  light  gray- 
ish vinaceous  or  vinaceous-buff.  The  rear  of  this  area  is  about  on  a  line  with 
front  line  of  arm  insertion  where  fold  comes  when  head  is  withdrawn.  When 
head  is  withdrawn,  the  fold  across  top  of  head  from  rear  of  one  eye  to  rear 
of  other  eye  is  prominent.  In  the  same  way,  another  fold  appears  on  the  venter 
just  in  front  of  arm  insertion.  Eye  is  like  that  of  female,  one  individual  having 
eye  with  reddish  cast,  other  having  green-yellow  eyes  like  the  female. 

Two  other  males  sent  by  H.  C.  Blanchard  from  Brownsville,  Tex.,  m 
August,  1931.  These  two  are  Saccardo's  olive  on  back.  The  bright  color  on 
legs  ranges  from  orange-cinnamon  to  ochraceous-salmon.  The  black  along 
the  side  of  one  is  a  heavy  unbroken  line,  of  the  other  a  band  of  broken  spots. 
The  stripe  from  vent  to  hind  legs  is  not  so  prominent  as  in  our  toads  from 
Kingsville. 

Structure:  Toes  and  fingers  slender,  feet  with  very  short  webs;  tubercles 
under  joints  of  toes  prominent;  inner  and  outer  sole  tubercles  large,  each  with 
a  cutting  edge;  legs  short;  short  fifth  toe  in  contrast  to  the  third  toe;  skin 
smooth,  loose,  and  leathery.  Fold  across  head  prominent  or  almost  lacking; 
eyes  prominent  and  beadlike. 


NARROW-MOUTH  TOADS:  BREVICIPITIDAE  565 

"Head  small,  body  large.  Limbs  short.  Muzzle  scarcely  longer  than  diameter 
of  eye,  projecting  a  little  beyond  mouth  border.  A  dermal  groove  across  head 
at  posterior  borders  of  eyelids,  and  one  from  below  posterior  canthus  of  eye 
to  shoulder.  Another  across  the  thorax  from  the  inferior  origin  of  one  humerus 
to  the  other.  Skin  everywhere  smooth.  Tympanic  drum  invisible.  When  the 
anterior  limb  is  extended  the  end  of  the  fore-arm  reaches  the  end  of  the 
muzzle.  The  distal  end  of  the  tarsus  reaches  the  anterior  base  of  the  humerus, 
and  the  end  of  the  second  toe  reaches  the  end  of  the  muzzle  when  the  hind 
limb  is  extended.  The  third  finger  is  rather  elongate,  and  the  lengths  of  the 
fingers  are  in  order,  beginning  with  the  shortest,  1-2-4-3,  &t  second  and 
fourth  being  equal.  In  the  posterior  foot  the  lengths  are,  beginning  with  the 
shortest,  1-2-5-3-4,  the  second  and  fifth  being  about  equal,  and  the  third  a 
good  deal  shorter  than  the  fourth.  The  palmar  tubercles  are  not  very  distinct. 
At  the  distal  end  of  the  tarsus  there  are  two  large  subequal  sharp-edged 
tuberosities.  The  edge  of  the  internal  is  oblique,  that  of  the  external  transverse. 
Distinct  small  tubercles  under  the  articulations  of  the  phalanges.  The  femur 
is  almost  entirely  inclosed  in  the  integument  of  the  body. 

"The  tongue  is  large,  and  forms  an  elongate  flat  ellipse.  The  interior  nos- 
trils are  anterior,  and  are  a  little  farther  apart  than  the  external  nostrils.  The 
latter  arc  nearly  terminal  in  position. 

14  Measurements  Meters 

Length  of  head  and  body  .041 

Length  of  head  to  rictus  oris  .006 

Length  to  axilla,  axially  .015 

Length  of  fore  limb  from  front  .022 

Length  of  fore  foot  .0095 

Length  of  hind  limb  from  ajius         .046 
Length  of  hind  foot  .023 

Width  of  head  at  rictus  oris  .010 

Width  of  extended  femora  .031 

Rather  abundant"  (E.  D.  Cope,  Tex.,  [1888]  1889,  p.  ?96). 

Voice:  "The  call  of  Hypopachus  cunetts  is  a  bleat  about  three  or  four  tones 
lower  than  that  of  Gastrophryne  olivacea  but  much  more  resonant.  Calls  last 
about  two  seconds  and  are  seldom  repeated  at  less  than  15  second  intervals, 
often  at  much  longer  intervals.  Whereas  Gastrophryne  prefers  to  support  it- 
self by  some  vertical  object,  Hypopachus  generally  floats  at  the  surface,  and 
from  these  positions  the  calls  are  made.  After  the  rains  cease  Hypopachus  be- 
comes silent  long  before  Gastrophryne"  (S.  Mulaik  and  D.  Sollberger,  Tex., 
1938,  p.  90). 

Breeding:  This  species  breeds  from  March  to  September  in  periods  of  heavy 
rain.  The  tadpole  is  small,  iM*2-i%  inches  (27-30  mm.),  wide,  and  flat,  like 
Microhyla.  It  has  a  broad  middorsal  band  of  grayish  olive,  with  scalloped 


566  HANDBOOK  OF  FROGS  AND  TOADS 

edges.  There  are  no  labial  teeth,  no  horny  mandibles,  no  papillae.  The  tad- 
poles transform  from  April  to  October,  at  %-%  inch  (10-12  mm.). 

June  16,  1930,  San  Benito,  Tex.  Went  to  the  breeding  pond  of  five  years 
ago.  Heard  no  frogs  and  took  a  very  few  tadpoles  with  hind  legs  well  de- 
veloped. With  early  heavy  rains  of  1930  the  species  probably  finished  by  June 
i  or  earlier. 

The  independent  shipments  of  live  Hypopachus  cuneus  in  August,  1931, 
by  C.  L.  Reed  and  H.  C.  Blanchard  might  imply  an  August  breeding  after 
heavy  rains.  The  females  from  Brownsville  were  gravid. 

We  secured  transformed  frogs  April  22,  1925,  at  San  Benito,  Tex.  They 
probably  transform  through  the  summer  and  fall.  Size,  10-12  mm. 

On  June  15, 1930,  at  the  same  pond  as  in  1925,  found  transformed  frogs  (37) 
which  measured  from  n.o  to  13.5  mm.,  one  16.0  mm.;  average,  12.3  mm., 
and  two  means,  12.0  mm.  and  12.5  mm. 

Notes  on  seven  of  these  transformed  frogs  follow: 

1.  16  mm.  This  was  the  only  one  of  the  37  that  had  a  light,  narrow  line  down 
middle  of  the  back.  This  same  specimen  had  a  light  median  line  on  the 
throat,  the  belly  blotched  with  white  spots,  and  fingers  and  toes  tessellated 
or  alternately  light-  and  dark-banded.  Pectoral  girdle  outlined  in  front  by 
black  line  which  goes  in  front  of  arm  to  eye. 

2.  13  mm.  Black  border  to  white  face-lme-to-be  goes  almost  to  meson  or  breast 
in  front  edge  of  pectoral  girdle  outline. 

3.  12.5  mm.  Black  line  from  eye  to  front  of  arm  to  pectoral  meson.  One  of  each 
side  meets  on  meson  in  a  V,  then  continues  caudad  on  meson  of  breast  and 
upper  belly  to  spiracle. 

4.  12  mm.  Tail  13  mm.,  tadpole  mouth,  white  beak.  Spiracle  scar  halfway  from 
pectoral  region  to  hind  legs. 

5.  12  mm.  Tail  15  mm.  Tail  tip  "purplish  black."  Looks  as  if  area  under  eye 
to  shoulder  is  white  or  gray. 

6.  12  mm.  Tadpole  mouth.  Beak  made  of  two  trough  ridges  ending  in  beak. 
Each  corner  of  mouth  with  clear  white  area  outlined  with  black.  Basal  tail 
musculature  clear  white  or  gray.  Tail  crests  heavily  blotched  with  black; 
tail  tip  or  caudal  half  of  tail  crests  and  musculature  almost  all  black. 

7.  12  mm.  Viewed  from  above  back  seems  to  have  two  white  areas  bordering 
dorsal  black  region,  which  extends  back  onto  tail,  with  musculature  on 
either  side  white. 

"The  senior  author  has  observed  the  eggs  being  laid  24  hours  after  the  be- 
ginning of  heavy  rains  from  April  to  October.  These  hatched  in  approxi- 
mately 12  hours  and  the  tadpoles  transformed  in  about  30  days.  The  gills  dis- 
appeared within  approximately  30  hours  after  hatching.  ...  A  complement 
of  approximately  700  eggs  was  taken  from  a  copulating  pair  of  Hypopachus 
cuneus  at  Edinburg,  Texas,  on  Sept.  24,  1935.  They  are  black  and  white  and 
measure  with  the  single  envelope  1.5  to  2  mm.  Without  the  envelope  they 


NARROW-MOUTH  TOADS:  BREVIClPlTlDAE  567 

measure  about  i  mm.  The  eggs  float  on  the  surface  of  temporary  pools  in 
rafts  loosely  held  together.  These  eggs  resemble  those  of  Gastrophryne  in  that 
the  envelope  is  truncate  and  they  float  with  this  flattened  surface  upward" 
(S.  Mulaik  and  D.  Sollberger,  Tex.,  1938,  p.  90). 

Journal  notes:  April  22, 1925,  San  Benito,  Tex.  One  mile  or  more  south  of 
town  at  Highland  found  on  east  edge  of  a  fine  blue  water-lily  pond  one 
Hypopachus  just  transformed.  It  was  in  the  wet  edges  where  B.  valliccps  trans- 
formed were  hopping  about.  Pseudacris  transformed  were  in  water's  edge. 
Found  two  or  three  more.  Found  no  mature  tads  in  this  pond;  must  be  species 
is  through  transforming  here.  One  had  a  long  tail  and  a  dark  dorsal  band 
with  concave  scalloped  edges.  In  a  fully  transformed  one  the  tail  stump  looks 
"bronzy,"  Under  parts  speckled  whitish  and  gray. 

June  15, 1930,  San  Benito,  Tex.  Same  pond,  opposite  Highland  School.  As  I 
approached  heard  a  note  I  thought  a  frog,  but  soon  saw  it  came  from  black- 
necked  stilts.  Found  in  the  grass  Pseudemys  elegans.  Does  it  feed  on  young 
Bufo  vatticeps  and  Hypopachus  cuneus?  Found  a  riband  snake  (T.  s.  proxi- 
mus).  When  I  stepped  on  it,  it  regurgitated  three  transformed  Hypopachus 
cuneus.  Then  I  made  it  give  up  more.  Had  nine  or  ten  of  these  little  narrow- 
mouthed  toads.  This  started  me  to  working  the  edge  of  the  pond.  Found  sev- 
eral transformed  ones  and  a  very  few  tadpoles.  A  few  Bufo  valliceps  trans- 
formed. Saw  another  riband  snake.  Made  it  disgorge.  One  of  these  narrow 
mouths  had  the  light  line  down  the  middle  of  the  back.  Others  do  not  have  it. 
No  cover  around  pond.  Snakes  collect  these  tiny  toads.  I  must  find  them;  once 
in  a  while  sec  a  transformed  H.  cuneus  in  grass  or  in  a  cow-punched  hole. 
There  being  no  apparent  cover,  in  my  desperation  I  resorted  to  turning  over 
dried  cow  clung.  Under  the  first  one  found  six  little  transformed  frogs.  Under 
an  old  shoe  found  several  Also  under  a  clump  of  matted  and  cut  weeds.  No 
adults  anywhere.  Waited  for  darkness.  None  heard. 

Authorities9  corner: 
E.  H.  Taylor  and  H.  M.  Smith,  Gen.,  1945,  p.  604. 


Genus  MICROHYLA  Tschudi 
(Formerly  Engystoma  and  Gastrophryne)  Map  37 

Recently  (April  i,  1946)  the  Hechts— Max  K.  and  Bessie  L.  Matalas— re- 
viewed this  group.  Their  conclusions  which  we  more  or  less  anticipated  are: 

1.  Microhyla  carolinensis  f  M.  olivacea,  and  M.  mazatlanensis  apparently 
interbreed  and  therefore  are  subspecies. 

2.  Microhyla  areolata  is  an  intermediate  between  carolinensis  and  olivacea. 

3.  The  Key  West  population  is  29  per  cent  olivacea-like,  48  per  cent  Key 
Westian,  and  23  per  cent  carolinensis-l ike— therefore  not  described  as  new. 

We  follow  the  old  pattern  yet  awhile  until  more  field  work  can  be  done 
even  though  we  agree  with  the  Hechts  that  M.  areolata  is  supported  by  very 
weak  evidence.  Their  paper  is  a  fine  paper,  conservative  on  Key  West,  and 
they  recognize  the  need  of  more  field  work  ("apparently  interbreed")  in 
their  first  conclusion. 

Mitchell's  Narrow-Mouth  Toad 

Microhyla  areolata  (Strecker).  Map  37. 

Range:  Southeastern  Texas.  Known  only  from  Victoria  and  Calhoun  coun- 
ties (}.  K.  Strecker,  Jr.,  Tex.,  1915,  p.  47).  More  recent  specimens  from  Houston 
to  western  Louisiana  have  been  so  identified  by  some  authors. 

Habitat:  Under  logs  and  similar  shelter. 

Size:  Adults,  %-i%  inches.  Males,  24-28  mm.  Females,  23-29  mm.  (From 
5  accessions  totaling  14  specimens  in  USNM.) 

General  appearance:  It  is  like  Microhyla  carolinensis,  but  the  back  is  areo- 
lated,  the  posterior  parts  even  pustular  (in  alcohol).  One  is  light  gray  with 
darker  markings  or  marblings,  which  are  heaviest  in  the  dorsal  region.  The 
limbs  are  heavily  marked.  Another  is  dark,  the  limbs  marked  with  blotches 
of  brownish  olive,  a  dark  line  from  orbit  to  orbit  along  the  muzzle,  and  a 
V-shaped  mark  between  the  orbits.  No  dark  line  extends  along  the  sides  as  in 
M.  carolinensis  (data  from  J.  K.  Strecker,  Jr.,  Tex.,  19093,  pp.  118-119). 

J.  K.  Strecker,  Jr.,  refers  the  following  notes  of  Dr.  Stejneger  to  this  form: 
"No.  35,942  (USNM)  Victoria,  Texas.  J.  D.  Mitchell,  collected  in  1897.  Skin 
of  back  areolated.  Four  specimens  (Victoria  High  School  Collection,  No.  52), 
collected  by  Mitchell  under  timber  in  Spring  Marsh,  Well  Camp,  Alligator 


NARROW-MOUTH  TOADS:  BREVICIPITIDAE  569 

Head,  Calhoun  County,  Texas,  March  1902,  also  have  the  back  areolated,  the 
posterior  parts  even  pustular.  Metatarsal  tubercles  rather  large  and  hind  feet 
short." 

Color  (in  alcohol) :  "Above,  light  gray,  with  darker  markings  which  are 
heaviest  in  the  dorsal  region.  Style  of  markings  might  be  termed  'marbling* 
on  account  of  their  irregular  outlines  and  light  colored  interspaces.  Limbs 
heavily  marked.  Under  surfaces,  light  gray  with  closely  placed  lightish  spots. 
Skin  of  back  areolated,  even  pustular  on  the  posterior  part.  Pustules  very 
uniformly  distributed.  .  .  .  Another  is  'rather  dark'  and  the  back  and  upper 
surface  of  the  limbs  are  marked  with  closely  placed  blotches  of  brownish  olive. 
In  the  type  a  dark  line  extends  along  the  muzzle  from  orbit  to  orbit  and 
there  is  a  small  dark  broad  V-shaped  mark  between  the  orbits.  No  dark  line 
along  the  sides  as  in  carolinense"  (J.  K.  Strecker,  Jr.,  Tex.,  1909,  pp.  118-119). 

Structure:  "Smaller  than  E.  carolinense.  .  .  .  Body  stout,  more  uniform  in 
width  than  E.  carolinense.  Muzzle  shorter  than  in  examples  of  E.  carolinense 
and  E.  texense  of  the  same  size.  Canthus  rostralis  not  prominent.  Hind  limbs 
short.  Hind  foot  unusually  short.  Inner  sole  tubercle  large.  ...  It  resembles 
E.  texense  in  size  and  in  the  shortness  of  the  hind  limbs  but  in  no  other  char- 
acteristic" (J.  K.  Strecker,  Jr.,  Tex.,  i909a,  p.  119). 

Because  of  its  restricted  distribution  and  the  presence  of  the  other  two 
species  in  the  same  place,  we  wonder  if  this  species  has  any  connection  with 
Dumeril  and  Bibron's  Engystoma  rugostim  (Parker,  1934,  suggests  M.  rugo- 
sum  as  a  possible  synonym),  which  Boulenger  reduced  to  G.  carolinensis. 
And  is  it  by  any  chance  the  roughened  "hotter  and  more  dry  season"  form 
which  Girard  intimates  might  exist  for  G.  olivacea?  Relative  measurements 
have  been  given  a  prominent  place  in  separating  the  three  species  of  Microhyla. 
We  find  these  differences  slight  and  append  here  a  table  of  a  few  measurements 
of  the  three  in  sizes  20  mm.  and  28  mm.  taken  from  specimens  in  the  National 
Museum,  and  one  set  from  a  frog  in  the  Cornell  University  collection. 

Name      carolinensis     olivacea      areo-       carolinensis  olivacea  arco- 

lata  lata 

No.  37063  37064  23744  38999  57688  48893  52296  1306  42334 
Sex  M  F  M  F  Type  M  F  F  M  M 

Length  20.0  20.0  20.0  20.0  20.0  28.0  28.0  28.0  28.0  28.0 
Head  to 

fold        5.5        5.5      5.0  5.0          5.0          6.0         4.8          6.5 

Head  to 

angle 

of 

mouth  5.0  4.5  4.0  5.0  4.0  5.0  5.5  5.0  5.0  5.5 
Snout  3.0  3.0  3.0  3.5  2.5  3.5  3.5  3.5  3-2  3-5 
Fore 

limb     n.o       9.5    11.0    9.0      10.0        12.0        11.0        11.5        11.5        12.5 


570  HANDBOOK  OF  FROGS  AND  TOADS 

Name     carolinemis     olivacea      areo-       carolincnsis  olivacca  areo- 

lata  lata 

Hind 

limb     26.0      23.5    23.0  22.5      21.0        29.0        29.0        31.0        27.0        26.0 
Hind 

foot 

with 

tarsus    14.5      13.0     13.0  13.5      12.0        16.0        17.0        16.5        14.0        15.5 

The  pustulate  or  areolate  character  of  M.  areolata  docs  not  impress  us  as 
noteworthy  when  we  consider  the  pustulate  character  in  the  posterior  region 
or  whole  dorsum  of  some  M.  carolinensis  specimens  in  the  Southeast.  In  fact, 
we  have  seen  there  some  individuals  more  pustular  than  any  specimen  of 
M.  areolata  revealed  to  date. 

Furthermore  the  types  and  some  of  the  other  specimens  were  of  young  or 
half-grown  individuals. 

Voice:  See  "Journal  notes"  and  "Authorities'  corner." 

Breeding:  See  "Journal  notes." 

Journal  notes:  June  13,  1925.  Arrived  at  Victoria  at  noon.  Rained  just  be- 
fore we  arrived.  Ought  to  be  Microhyla  areolata  in  railroad  ditches.  No  frog 
notes. 

June  14.  Went  along  railroad  right  of  way.  Turned  over  cover.  Saw  Cnemi- 
dophorus  but  no  A/,  areolata,  which  we  consider  a  debatable  form. 

April  17,  1934,  Spring  Creek  near  Houston,  Tex.  Prof.  A.  C.  Chandler  of 
Rice  Institute  took  us  to  the  bottom  lands  where  he  and  Harwood  first  found 
the  Microhyla  which  they  considered  areolata.  We  searched  under  and  in  logs 
but  did  not  succeed  in  catching  any.  Much  rain  came  last  night  and  on  days 
before,  and  soon  in  a  shady  overflow  of  the  woods  we  heard  a  chorus  of 
narrow-mouths.  We  also  heard  a  few  Rana  catesbeiana.  Many  R.  clamitans, 
a  few  Acris,  and  no  end  of  Microhyla.  At  the  base  of  a  tree  would  be  drift- 
wood and  a  mass  of  needles.  In  these  masses,  usually  beneath,  we  would  hear 
the  little  chorister,  but  we  could  never  locate  one  and  we  had  no  net.  The 
note  sounds  like  that  of  M.  carolinensis.  Later,  beside  the  edge  of  the  overflow 
in  trash  Anna  found  little  packets  of  5-12  eggs.  They  were  very  scattered. 
Either  the  pair  moved  or  the  masses  had  broken.  These  small  masses  suggest 
M.  texensis  (olwacea)  in  packet  size  and  inconspicuousness  of  eggs.  There 
were  many  of  these  in  the  dumbbell  form  of  development. 

April  18.  Stopped  at  W,  A.  Bevans'  reptile  garden,  Houston,  Tex.  Found 
he  had  a  Microhyla  in  the  snake  cage  for  food.  He  gave  it  to  me.  He  said  they 
were  common  around  the  reptile  house  and  he  feeds  these  frogs  to  his  coral 
snakes  and  other  snakes. 

Authorities'  corner:  "In  Texas  ...  I  heard  it  in  the  streets  of  Houston  and 
San  Antonio.  In  the  former  city  it  was  abundant,  in  copula,  in  the  ditches  that 


NARROW-MOUTH  TOADS:  BREVICIPITIDAE 


571 


border  some  of  ihe  streets,  in  September.  The  cry  is  loud  for  the  size  of  the 
animal,  and  is  similar  to  that  of  Bufo  americanus,  except  in  being  higher 
pitched  and  more  nasal  (in  the  vulgar  sense).  The  animals  are  extremely  shy, 
and  become  silent  on  the  approach  of  human  footsteps;  and  as  only  the  tip 
of  their  nose  projects  above  the  water-level  they  disappear  beneath  it  without 
leaving  a  ripple"  (E.  D.  Cope,  Gen.,  1889,  p.  386). 

"June  15,  1918.  A  heavy  thunder  shower  brought  out  the  Narrow-mouthed 
Toads.  Found  them  abundant  and  noisy  in  a  large  rain  pool  about  ten  o'clock 
at  night.  The  call  is  harsh  and  grating,  almost  a  bleat.  It  is  repeated  frequently, 
but  not  as  rapidly  as  that  of  the  Hylas.  The  males  sit  in  the  water  in  the  thick 
grass  that  fringes  the  pools,  and  are  almost  impossible  to  find  with  a  light, 
although  they  are  not  at  all  shy.  Collected  six  specimens  by  treading  down 
the  grass  into  the  water  and  catching  them  as  they  swam.  They  swim  slowly 
with  their  short  legs  and  webless  feet  and  do  not  dive  when  disturbed  as 
frogs  and  toads  do.  From  this  date  until  the  middle  of  August  I  heard  them 
frequently,  but  seldom  in  any  abundance.  After  a  rain  could  be  heard  now 
and  then  in  a  pool  or  roadside  ditch  before  sunset  as  well  as  at  night"  (P.  H. 
Pope,  Tex.,  1919,  p.  94). 

Frankly  we  never  have  believed  very  strongly  in  this  form.  We  have  re- 
tained it  partly  because  of  attachment  for  its  author  and  partly  because  this 
volume  should  be  conservative  and  suggestive. 

Narrow-Mouth  Toad,  Narrow-Mouth  Frog,  Narrow-mouthed  Frog, 
Narrow-mouthed  Toad,  Nebulous  Toad,  Toothless  Frog 

Microhyla  carolinensis  (Holbrook).  Plate  CXXV;  Map  37. 

Range:  Maryland  to  Florida  to  Texas.  North  to  Nebraska,  Illinois,  southern 
Indiana,  and  Kentucky. 

Habitat:  Nocturnal,  a  lover  of  rain,  cover,  and  moist  situations.  It  is  a  sub- 
terranean species.  Hides  in  or  beneath  decaying  logs  or  under  haycocks  or 
other  shelter. 

One  would  never  find  it  during  the  day  if  he  did  not  hear  it  bleat  during  a 
rainstorm  or  cloudy  weather,  or  if  he  did  not  unearth  it  from  beneath  its 
cover.  They  may  appear  by  day  during  a  drenching  rain.  Sometimes  the  din  is 
incredible.  They  apparently  do  not  have  far  to  migrate  when  the  call  of  the 
rain  comes. 

"Found  under  rocks,  boards,  and  debris;  moderately  common"  (O.  C. 
Van  Hyning,  Fla.,  1933,  p.  4). 

"Apparently  this  species  has  not  been  recorded  from  the  state.  We  heard 
several  calling  from  thick  grass  in  a  cattail  swamp  a  few  hundred  feet  from 
Cove  Point  lighthouse.  The  water  was  slightly  brackish  but  Hyla  cinerea  and 
H.  versicolor  were  present.  Several  other  Gastrophryne  were  collected  in  a 
fresh  water  swamp  between  Cove  Point  and  Solomon's  Island.  These  were 


572  HANDBOOK  OF  FROGS  AND  TOADS 

also  calling  from  thick  tussocks  of  grass  with  usually  the  rear  part  of  the  body 
submerged.  The  call  seemed  to  us  to  resemble  that  of  Bufo  jowleri  but  was 
more  explosive,  shorter  and  higher  pitched"  (G.  K.  Noble  and  W.  G.  Hassler, 
Md.,  1936,  p.  63). 

"Males  were  heard  calling  on  May  8  at  the  time  of  the  earliest  spring  ap- 
pearance. The  call  resembles  the  bleat  of  a  kid.  They  appear  to  spend  much 
time  in  the  vicinity  of  the  pools  probably  waiting  favorable  conditions  of 
high  humidity  and  heavy  rains  to  begin  breeding.  On  July  3  many  were 
found  hidden  under  pine  needles  in  the  vicinity  of  'low  ground.'  After  heavy 
rains  in  early  August  they  were  in  the  puddles  and  ponds,  calling  and  clasping 
in  tremendous  numbers.  They  utilized  the  edges  of  permanent  ponds  as  well 
as  the  most  temporary  puddles.  Active  individuals  were  taken  on  the  night  of 
November  6"  (13.  B.  Brandt,  N.C.,  1936,  p.  222). 

"Little  affected  by  major  habitat  boundaries;  under  rocks,  logs,  and  piles 
of  debris;  usually,  but  not  always,  in  moist  soil.  Abundance. — Solitary  and 
secretive,  but  common  in  all  parts  of  its  Florida  range.  The  breeding  choruses 
are  sometimes  very  large.  Habits. — Lucifugous;  I  have  never  seen  one  in  the 
open  in  the  daytime.  Despite  its  rotund  belly  and  short  legs  Microhyla  is  nim- 
ble and  active  when  frightened.  I  have  seen  several  dive  into  the  mouths  of 
crayfish  burrows  when  the  logs  under  which  they  were  hiding  were  over- 
turned; in  loose  leaf -mold  they  rapidly  burrow  out  of  sight.  In  Key  West 
they  occur  in  old  lumber  and  trash  piles  with  the  geckos  and  skinks"  (A.  F. 
Carr,  Jr.,  Fla.,  1940,  p.  68). 

Size:  Adults,  %-i%  inches.  Males,  20-34  mm.  Females,  22-36  mm. 

General  appearance:  This  frog  is  small,  dark-colored,  and  relatively  smooth- 
skinned.  A  fold  of  skin  extends  across  the  head  just  back  of  the  eye.  The  head 
is  small,  the  snout  pointed.  As  one  child  remarked,  its  head  looks  like  a  tiny 
turtle  head.  The  fold  of  skin  accentuates  this.  There  is  no  tympanum.  The 
hind  limbs  are  proportionally  stout.  The  fingers  and  toes  are  without  webs. 
The  back  may  be  black,  brown,  or  gray,  the  under  parts  dusky  gray  or  brown 
speckled  or  mottled  with  light. 

Color:  Male.  Top  of  head  and  upper  eyelids  mineral  gray,  pale  smoke  gray, 
or  with  yellowish  cast.  Back  dusky  with  fine  bluish  white  spots  or  dots.  Under 
parts  dusky  and  bluish  white.  Throat  with  fine  dots  not  blotches,  therefore 
looks  blacker  in  male.  Pupil  circular. 

Female.  Lighter,  larger.  Practically  no  spots  on  pectoral  region.  None  on 
throat.  Throat  same  color  as  belly.  Practically  no  rusty  spots.  Ground  color 
of  the  dorsum  more  greenish,  i.e.,  olive-gray,  mineral  gray,  or  grayish  olive.. 
Sometimes  at  breeding  season  males  and  females  may  be  alike  in  color;  for 
instance,  on  April  24,  1921,  several  pairs  were  alike,  some  reddish,  one  gray. 
Most  of  the  pairs,  however,  were  diverse. 

Structure:  Skin  usually  smooth  or  finely  tuberculate;  upper  jaw  projecting 
beyond  the  lower;  a  fold  of  skin  extending  from  one  arm  insertion  to  eyes, 


NARROW-MOUTH  TOADS:  BREVICIPITIDAE  573 

thence  to  other  arm  insertion;  tongue  broad,  only  caudal  border  free;  inner 
metatarsal  tubercle  present. 

Voice:  The  call  is  a  bleating  baa  that  might  well  deceive  anyone.  They 
usually  call  from  the  water  with  rear  parts  submerged  and  fore  feet  planted 
on  the  bank  or  on  some  other  support  such  as  trash  on  the  water's  surface.  Oc- 
casionally they  are  out  on  the  bank  or  in  a  grassy  tussock.  The  throat  swells 
out  like  the  light  bubble  of  a  toad  or  spadefoot.  The  call  lasts  1.0-2.0  seconds. 
Though  he  had  never  caught  the  frog  and  did  not  know  which  one  it  was, 
Le  Conte  (Gen.,  1856,  p.  431)  gave  a  very  good  description  of  Microhyla's 
call:  "There  is  in  the  waters  of  ponds  and  ditches  a  small  frog  whose  note 
exactly  resembles  the  bleating  of  a  lamb,  so  truly  as  to  deceive  anyone."  As  a 
general  rule,  one  must  be  within  20-60  yards  to  hear  Microhyla's  baaa.  At 
times  the  male  has  one  or  two  other  notes  not  of  this  order.  The  frog  becomes 
vertical  or  bent  backward  with  the  effort  of  calling.  On  May  21,  1921,  F. 
Harper  and  I  counted  the  intervals  and  periods  of  calls.  The  record  of  thirty- 
seven  intervals  ranged  from  3-58  seconds,  at  least  20  lasting  30  seconds  or 
more.  The  actual  call  in  most  instances  occupied  1.5-2.0  seconds.  The  intervals 
of  another  calling  frog  were  from  1-60  seconds  with  only  one  below  10  seconds. 

Breeding:  This  species  breeds  from  May  1  to  Sept.  1.  The  egg  mass  is  a 
surface  film,  or  smaller  packets  of  10-90  eggs;  the  complement  is  850  eggs. 
The  black  and  white  eggs  are  firm  and  distinct  like  glass  marbles,  making  a 
fine  mosaic;  the  envelope  %-/{;  inch  (2.8-4.0  mm.)  is  a  truncated  sphere,  flat 
above.  The  egg  is  ^.v/^o  inch  (r. 0-1.2  mm.).  The  tadpole  is  small,  i  inch 
(26.5  mm.)  flat,  wide,  and  elliptical  in  shape;  the  snout  is  truncate;  the  dorsal 
and  ventral  sides  of  the  head  are  flattened;  and  the  eyes  are  visible  from  the 
ventral  aspect.  The  tail  is  medium,  obtuse  or  rounded,  sometimes  with  a 
black  tip.  The  spiracle  is  median,  closely  associated  with  the  anus,  and  its  sepa- 
ration apparent  as  the  hind  legs  appear.  There  arc  no  teeth,  no  horny  mandi- 
bles, no  papillae.  After  a  period  of  20  to  70  days,  the  tadpoles  transform  from 
mid-June  to  mid-October,  at  %G-%  inch  (8.5-12  mm.). 

"April  i  to  September  3.  They  breed  during  or  after  heavy  rains  in  puddles 
or  the  inundated  margins  of  ponds  and  ditches.  The  singing  males  usually 
conceal  themselves  in  grass  or  trash  at  the  water's  edge.  On  July  14,  1935, 
I  saw  a  small  chorus  in  a  rain  puddle  in  bare  white  sand  on  the  banks  of 
Weekiwachee  Spring  Run;  here  the  males  had  all  wriggled  their  bodies  into 
the  wet  sand  and  were  calling  with  only  the  snouts  protruding;  this  puddle 
could  not  possibly  have  lasted  more  than  a  few  hours  after  the  cessation  of  the 
rain"  (A.  F.  Carr,  Jr.,  Fla.,  1940,  p.  68) . 

"A  nocturnal  species  very  hard  to  find  except  in  the  breeding  season,  which 
may  be  exemplified  by  the  fact  that  although  I  caught  forty  the  first  time  I 
ever  found  them  breeding,  I  have  only  seen  two  or  three  at  Raleigh  apart  from 
the  breeding  season. 

"The  breeding  season  is  from  May  to  August  and  these  frogs  seem  to  prefer 


574  HANDBOOK  OF  FROGS  AND  TOADS 

a  night  when  a  warm  drizzle  is  falling.  The  cry  is  loud,  easy  to  recognize  but 
hard  to  describe,  and  the  only  description  I  have  ever  seen  that  seemed  to  fit 
was  the  one  that  compared  it  to  an  electric  buzzer.  The  tadpoles  are  dark 
colored  and  transform  to  tiny  frogs  the  same  season"  (C.  S.  Brimley,  N.C., 
1941,  p.  3). 

Journal  notes:  May  21, 1921,  Okefinokee  Swamp.  Went  to  old  and  new  hog 
hole  and  heard  a  Microhyla,  a  bleat  like  a  lamb  or  calf.  Came  home  pell-mell 
and  heard  Microhyla  in  ditch  in  Negro  quarters.  We  started  at  8  P.M.  Air  70°?. 
We  stopped  to  find  them  in  ditch.  Throat  like  Btifo  throat,  black  in  color. 

May  22,  Okefinokee  Swamp,  Ga.  In  the  ditches  were  Microhyla  eggs.  Each 
egg  stands  out  distinctly.  We  found  the  water  blackish,  trashy,  and  oily  with 
floating  packets  of  eggs.  Some  masses  were  i  by  i  inch  (25  by  25  mm.)  in 
diameter;  others  2  by  i  inches  (50  by  25  mm.) ;  others  2  by  2  inches  (50  by 
50  mm.) ;  some  round  masses,  others  square.  Each  jelly  envelope  abuts  that  of 
the  next  in  pentagonal  or  hexagonal  fashion.  In  proper  light  a  mass  of  eggs 
makes  a  mosaic.  Some  egg  masses  along  banks  in  weeds  may  be  i  foot  10 
inches  (550  mm.)  long  and  3  or  4  inches  (75  or  100  mm.)  wide.  One  mass 
in  the  middle  of  ditch  was  among  chips  and  was  4  by  7  inches  (100  by  175 
mm.).  There  are  few  such  in  mid-pond.  We  believe  they  lay  large  masses 
along  the  edges  or  among  the  brush,  and  the  wind  scatters  them. 

In  a  clean  pond  (20  by  3  feet)  along  its  edges  among  grass  was  one  packet 
of  eggs,  100-125.  They  are  brown  and  yellowish,  more  eggs  in  a  mass  than  in 
a  film  of  Hyla  versicolor.  Insects  get  into  the  fresh  masses. 

Authorities9  corner: 

J.  E.  Holbrook,  Gen.,  1842,  p.  24.  E.  Loennberg,  Fla.,  1895,  p.  338. 

J.  A.  Ryder,  N.C.,  1891,  p.  838.  R.  Barbour,  Ky.,  1941,  p.  262. 

Taylor's  Microhyla 

Microhyla  mazatlanensis  Taylor.  Map  37. 

Range:  "The  third  race,  Microhyla  c.  mazatlanensis,  probably  is  generally 
distributed  in  the  foothills  of  northwestern  Mexico  (Sinaloa  and  Sonora) 
barely  ranging  into  southern  Arizona;  roughly  its  range  may  be  said  to  oc- 
cupy the  eastern  portion  of  the  Sonoran  and  Sinaloan  biotic  provinces^  as 
mapped  by  Dice.  ...  If  our  belief  concerning  the  identity  of  the  type  is  cor- 
rect, mazatlanensis  is  known  from  the  following  localities:  Mazatlan,  Sinaloa 
(type  locality) ;  10  miles  north  of  Pilares,  5  miles  north  of  Noria,  and  from 
Guirocoba  in  Sonora;  and  Pefia  Blanca  Springs,  in  Santa  Cruz  County,  Ari- 
zona" (M.  K.  Hecht  and  B.  L.  Matalas,  Gen.,  1946,  pp.  5-6). 

Habitat:  A  lowland  form;  under  rocks,  in  piles  of  stones,  at  edge  of  and  in 
pools,  boggy  areas,  and  overflow  areas. 

Size:  Adults  smaller  than  M.  olivacea,  %-iV^  inches.  Males,  22-28  mm.  Fe- 
males, 22-30  mm. 


NARROW-MOUTH  TOADS:  BREVICIPITIDAE  575 

General  appearance:  "Toes  without  webs,  the  outer  metatarsal  tubercle 
absent.  Related  to  Mtcrohyla  olivacea  but  distinctly  smaller  in  size,  the  head 
a  little  narrower,  the  snout  projecting  more,  a  little  more  flattened  above  and 
rounding  at  the  tip;  eye  smaller  proportionally;  choanae  smaller,  largely  con- 
cealed by  the  shelf  from  the  maxilla  when  seen  from  below;  toes  and  fingers 
more  rounded  without  a  lateral  ridge.  An  indistinct  black  stripe  behind  eye 
to  some  distance  on  side.  Brown  above"  (E.  H.  Taylor,  Ariz.,  1943,  p.  355). 

"From  the  diagnosis  given  little  can  be  found  that  warrants  the  description 
of  a  new  form  although  the  form  appears  to  be  recognizable  on  the  basis  of 
pattern  characters.  .  .  .  Presumably  the  series  is  composed  of  small  adults, 
and  it  is  for  this  reason  that  the  head  appears  to  be  narrower  and  the  snout 
longer.  It  has  been  observed  that  smaller  specimens  of  both  carolinensis  and 
olivacea  tend  to  have  more  acute  snouts  and  relatively  narrower  heads  than 
adults  approaching  full  maturity  and  maximum  size.  Perhaps  the  most  use- 
ful data  given  are  those  in  the  color  pattern"  (M.  K.  Hecht  and  B.  L.  Matalas, 
Gen.,  1946,  p.  4). 

"Ventrum  immaculate  or  with  scattered  melanophores.  A  blotch  or  spots 
on  the  femur  and  tibia  which  form  a  bar  or  continuous  line  when  the  limb 
is  folded;  dorsum  with  dark  spots.  Mazatlanensts"  (M.  K.  Hecht  and  B.  L. 
Matalas,  Gen.,  1946,  p.  7). 

Structure:  (Omitted.  Sec  "General  Appearance,"  "Color,"  and  "Authorities' 
corner.") 

Color:  "Above  nearly  uniform  brown  with  some  black  spots  tending  to 
form  a  pattern  medially;  a  broken  black  line  begins  behind  the  eye  and  con- 
tinues to  some  distance  on  the  side;  trace  of  an  inguinal  spot,  which  together 
with  a  single  bar  or  spot  on  the  femur,  the  tibia  and  foot,  form  a  continuous 
line  when  the  limb  is  folded.  Venter  cream,  with  a  very  slight  peppering  of 
pigment  on  the  chin;  sides  slightly  mottled  with  lighter  and  darker;  under- 
side of  the  feet  purplish  brown,  and  the  hands  similarly  colored,  but  to  a 
lesser  extent"  (E.  H.  Taylor,  1938,  p.  356). 

"Taylor's  mazatlanensis  is  virtually  identical  with  the  specimens  from  Pena 
Blanca  Springs,  Santa  Cruz  County,  Arizona  (U.  M.  M.  Z.  Nos.  75737,  75738, 
totaling  15  specimens),  and  those  from  Sonora  10  miles  north  of  Pilares 
(U.  M.  M.  Z.  No.  78333),  Guirocoba  (A.  M.  N.  H.  Nos.  51244-51247)  and 
Noria  (U.  M.  M.  Z.  Nos.  72174-72177)  which  have  these  same  markings  .  .  ." 
(M.  K.  Hecht  and  B.  L.  Matalas,  Gen.,  1946,  p.  4). 

Voice  and  Breeding:  Our  only  evidence  comes  from  B.  Campbell's  first 
collection  of  this  form  and  from  J.  W.  Hilton's  notes  for  Bogert  and  Oliver. 

Considering  it  Gastrophryne  olivacea  (Hallowell)  B.  Campbell  wrote: 

"Adults:  Pena  Blanca  Springs,  15,  July  11  and  20.  Larvae:  Pena  Blanca 
Springs,  Aug.  2.  After  one  of  the  first  heavy  rains  of  the  season  at  Pena  Blanca 
Springs,  we  heard  strange  noises  coming  from  a  boggy  spot  at  the  overflow 
of  the  reservoir.  We  stalked  the  source  with  a  flashlight  and  were  surprised  to 


576  HANDBOOK  OF  FROGS  AND  TOADS 

find  narrow-mouth  toads,  which  have  been  reported  but  once  (Sonora,  Mex- 
ico) west  of  Texas.  They  were  squatting  in  the  half  inch  or  so  of  water  be- 
tween the  hillocks  of  grass  and  calling  at  approximately  twelve-second  inter- 
vals, mostly  in  unison.  One  was  heard  from  a  pool  down  the  canyon  but  was 
not  collected. 

"On  the  evening  of  July  20,  we  went  up  the  canyon  to  investigate  a  chorus 
of  Hyla  arenicolor  which  we  had  heard  for  several  nights.  A  number  of 
Gastrophryne  were  found  in  the  pools,  croaking.  Curiously  enough,  they 
seemed  to  be  limited  to  the  pools  at  the  bases  of  trees;  possibly  they  remained 
among  the  roots  in  the  daytime.  Several  pairs  were  found  in  amplexation, 
both  in  the  pools  and  just  out  of  the  water.  The  males  have  round  blackish 
voice  pouches  which  swell,  in  croaking,  to  the  size  of  a  small  marble.  Some 
males  were  sprawled  out  on  the  water  after  the  manner  of  Scaphiopus,  but 
most  of  them  were  standing  upright  in  the  water  next  to  the  roots  of  trees 
or  near  the  bases  of  large  rocks,  with  their  heads  emerging.  These  toads  were 
heard  in  a  small  pool  in  a  gully  near  the  top  of  the  canyon  wall  on  the  evening 
of  July  22.  About  seven  miles  east  and  somewhat  north  of  Patagonia,  at  a 
little  station  named  Vaughan,  I  heard  them  croaking  on  the  evening  of 
August  23. 

"We  found  no  eggs  of  this  species,  but  collected  the  larvae  from  the  canyon 
pools  at  Pena  Blanca  Springs"  (B.  Campbell,  Ariz.,  1933,  pp.  6-7). 

"Three  males  and  one  mature  female,  with  well  developed  eggs,  were 
collected  by  Hilton  [at  Guirocoba,  Sonora].  The  largest  male  measures  28.7 
mm.  in  snout-vent  length;  the  female  measures  29.6  mm. 

"Both  Allen  ([Ariz.,]  1933,  p.  3)  and  Taylor  (i938b,  p.  516  [Gen.  i936aj; 
I94ob,  p.  531)  have  suggested  the  possibility  that  Sonoran  and  Sinaloan  speci- 
mens represent  an  unrecognized  form.  Sufficient  material  is  not  at  hand  to 
verify  this. 

"Hilton  says  in  his  field  notes,  'This  is  one  of  the  noisiest  amphibians  in 
the  arroyos  but  difficult  to  locate.  They  sit  under  dead  leaves  and  twigs  along 
the  bank  and  make  a  shrill  call  that  sounds  more  like  that  of  a  bird.  This  frog 
is  called  "rana  Pajanto"  and  is  not  easy  to  capture' "  (C,  M.  Bogert  and  J.  A. 
Oliver,  Gen.,  1945,  pp.  343-344)- 

Journal  notes:  The  three  times  we  have  been  at  Pena  Blanca  Springs  have 
been  of  short  duration  and  not  in  a  rainy  season,  and  we  therefore  wholly 
missed  this  form. 

Authorities9  corner:  The  first  record  of  this  form  came  in  Allen's  capture 
of  four  at  Noria,  Sonora. 

"The  specimens  are  all  males,  with  a  head  and  body  length  of  25  to  30  mm. 
Further  collecting  in  Mexico  will  be  necessary  before  the  status  of  this  Gas- 
trophryne is  clearly  understood.  It  differs  distinctly  from  G.  usta  in  colora- 
tion and  in  having  only  one  metatarsal  tubercle  and  from  elegans  in  coloration 
and  proportions.  The  coloration  is  that  of  texensis,  and  the  proportions  are 


NARROW-MOUTH  TOADS:  BREVICIPITIDAE 


577 


about  the  same,  though  slight  differences  not  evident  in  proportional  measure- 
ments seem  apparent  to  the  eye.  The  Sonoran  specimens  will  have  to  stand 
as  texensis  for  the  present,  in  spite  of  the  wide  gap  in  the  known  range.  Found 
about  the  same  pools  as  Pternohyla  jodiens"  (M.  J.  Allen,  Ariz,  and  Sonora, 

*933>  P-  3)- 

In  July  15, 1936,  E.  H.  Taylor  gave  us  his  first  record  of  this  species :  "Micro- 
hyla  olivacea.  ...  I  have  tentatively  associated  with  this  species  three  small 
microhylids  (Nos.  1236  to  1238)  collected  about  two  miles  east  of  Mazatlan 
under  rocks  at  the  base  of  a  small  clay  hill.  When  compared  with  Texas  speci- 
mens of  equal  size  they  differ  in  having  a  narrower  head,  the  snout  a  little 
more  projecting  and  more  flattened.  They  are  somewhat  darker  and  on  the 
side,  from  snout  to  groin,  the  pigment  tends  to  form  a  darker  broken  line. 
There  is  a  slight  difference  in  the  shape  of  the  foot  and  the  metatarsal  tubercle 
is  slightly  more  salient.  They  differ  somewhat  less  from  a  series  of  specimens 
collected  by  Hobart  Smith  and  David  Dunkle  at  Conefos,  Durango.  None  of 
the  three  specimens  approaches  the  maximum  si/e  of  olivacea.  A  larger,  more 
representative  series  may  demonstrate  that  these  and  perchance  other  char- 
acters warrant  a  specific  designation  for  the  coastal  form.  If  properly  associated 
with  olivacea  these  records  extend  the  known  range  some  200  miles  farther 
to  the  southwest"  (E.  H.  Taylor,  Gen.,  1936^  p.  516). 

In  1943  Taylor  in  describing  this  form  as  a  new  species  remarked :  "These 
small  specimens  were  obtained  from  a  pile  of  rocks  near  a  small  temporary 
rain  pool.  In  the  same  pile  of  stones  I  obtained  the  type  of  Bufo  mazatlanensis. 

"These  specimens  were  originally  associated  with  Microhyla  olivacea,  al- 
though the  differences  were  recognized  at  that  time.  .  .  .  Since  then  I  have 
examined  large  series  of  this  genus  from  Mexico  and  have  a  better  idea  of  the 
amount  of  variation  that  may  be  expected.  This  has  caused  me  to  regard  this 
lowland  form  as  distinct  from  A/,  olivacea. 

"The  species  differs  from  Microhvla  elcgans,  in  lacking  the  heavy  pigmenta- 
tion of  the  ventral  surfaces,  and  the  broad  lateral  stripe  which  begins  on  the 
tip  of  the  snout.  Elegans  has  well-developed  lateral  ridges  or  fringes  on  the 
sides  of  most  of  the  digits,  and  the  tips  are  flattened  and  somewhat  widened. 
The  inguinal  spot  is  strongly  defined. 

"Microhyla  usta  has  been  reported  from  Presidio  in  the  southern  part  of 
Sinaloa  and  may  also  occur  in  this  locality.  This  may  be  separated  easily  by 
the  fact  that  usta  has  a  very  large  inner  metatarsal  tubercle,  and  also  a  strongly 
developed  outer  one,  while  the  described  form  has  a  small  inner  and  no  outer" 
(E.  H.  Taylor,  Gen.,  1943,  p.  357) . 


578  HANDBOOK  OF  FROGS  AND  TOADS 

Texas  Narrow-Mouth  Toad,  Texas  Toothless  Frog, 
Western  Narrow-mouthed  Frog 

Microhyla  olwacea  (Hallowell).  Plate  CXXVI;  Map  37. 

Range:  Western  Missouri  and  eastern  Nebraska  and  Kansas  through  Okla- 
homa and  central  Texas  to  Coahuila  (Taylor-Smith,  Schmidt  and  Owens), 
Chihuahua  (Taylor  and  Smith),  and  Durango  (Smith  and  Dunkle), 

Habitat:  Nocturnal  in  habit.  Seek  protection  under  logs  or  dead  stumps 
sunken  in  the  ground;  in  the  far  southern  tip  of  Texas  under  fallen  trunks  of 
Spanish  bayonet.  Breed  in  ponds,  roadside  ditches,  or  temporary  rain  pools. 

"Fairly  common  in  damp  soil,  under  pieces  of  limestone  or  sandstone.  The 
earliest  spring  date  is  April  i"  (E.  R.  Force,  Okla.,  1930,  p.  27). 

"As  Dice  (1923),  who  found  this  species  on  the  Rocky-ground  community 
and  Hillside  forest  community  of  Riley  county,  indicates,  this  species  of  frog 
in  Kansas  is  far  more  characteristic  of  terrestrial  than  of  aquatic  habitats.  Of 
the  numerous  specimens  I  have  collected  or  seen  collected  near  Manhattan, 
all  were  from  communities  mentioned  by  Dice.  .  .  .  The  usual  cover  is  rocks" 
(H.  M.  Smith,  Kans.,  1934,  p.  503). 

"The  Texas  narrow-mouthed  toad  is  common  and  numerous  in  all  parts 
of  Oklahoma.  It  is  especially  abundant  in  the  prairies  and  in  the  oak-hickory 
savannah  areas*'  (A.  N.  Bragg,  Okla.,  19413,  p.  52). 

Size:  Adults,  %-i%  inches.  Males,  20-33  mm.  Females,  19-38  mm. 

General  appearance:  This  is  a  small,  usually  dark-colored,  smooth-skinned 
"frog-toad"  with  small,  pointed,  flattened  head.  The  under  parts  are  uni- 
formly white,  the  head  and  body  depressed,  the  limbs  slender  in  appearance. 
The  back  has  black  spots.  The  eyes  are  small  and  beadlike. 

Color:  March  27,  1925,  Beeville,  Tex.  Male.  Grayish  olive,  or  citrine-drab 
on  back.  Spots  on  back  black.  Color  on  side  wood  brown,  buffy  brown,  or 
olive-brown.  Rear  of  thighs  with  some  spots  of  the  three  browns  above.  Throat 
violet-slate,  rim  of  lower  jaw  like  back  or  sides.  Pectoral  region  and  front 
half  of  breast  whitish  with  seafoam  green  cast.  Abdominal  region  and  under- 
side of  thighs  with  some  pinkish  vinaceous  cast.  Pupil  round.  Iris  spotted, 
grayish  olive  or  yellow-green  with  wood  brown  or  other  browns.  Fingers 
cartridge  buff.  Foot  like  upper  parts  of  limbs  but  with  whitish  tips. 

Female.  Throat  light.  Fingers,  forefeet  white.  Underside  of  hind  feet  gray- 
ish. Upper  parts  grayish  olive  or  light  grayish  olive.  Irregular  lateral  stripe  of 
light  vinaceous-drab  or  brownish  drab. 

Structure:  Body  more  depressed  than  in  M.  carolinensis  and  head  appearing 
more  pointed,  because  of  depressed  body;  limbs  more  slender  in  M,  olivacea. 

Three  characters  often  emphasized  prove  neither  constant  nor  absolute 
points  of  difference  between  the  two  species. 

i.  Girard  held  that  the  cleft  of  the  mouth  does  not  extend  as  far  back  in 


NARROW-MOUTH  TOADS:  BREVICIPITIDAE 


579 


%&£'  C3- '?;S:  :*IIP ' 'S 


CXXV.  Microhyla  carolinensis 
XO-  1,2,6,7.  Females.  3.  Male  croaking. 
.  Tadpole.  5.  Egg  mass. 


Plate  CXXVL  Microhyla  olivacea.  1,7. 
Females  (X^/0-  2,6.  Males  (X^)-  3- 
Eggs  (X%)-  4^5-  Transforming  frogs 

(Xi). 


580  HANDBOOK  OF  FROGS  AND  TOADS 

M.  olivacea  as  in  M .  carolinensis.  Sometimes  specimens  of  exactly  the  same 
length  have  it  farther  back  in  M .  olivacea  than  in  M.  carolinensis. 

2.  Miss  Dickerson  stated  that  the  head  to  shoulder  length  is  one-third  the 
total  length  in  M.  olivacea  and  one-fourth  in  M.  carolinensis.  Some  relative 
measurements  of  M.  olivacea  from  Beeville,  Tex.,  and  M.  carolinensis  from 
Okefinokee  Swamp,  Ga.,  are  as  follows : 

HEAD  TO  SHOULDER  CONTAINED  IN  LENGTH 

M.  olivacea  M.  carolinensis 

Males  2.4          2.5  2.3  2.6 

Females  2.8          2.95  2.5          2.9 

The  difference  is  much  less  than  was  stated  by  Miss  Dickerson. 

3.  Strecker  holds  that  the  greatest  width  of  body  in  M.  olivacea  is  usually 
less  than  one-half  the  length.  A  few  random  comparisons  of  the  same  Bee- 
ville and  Okefinokee  Swamp  materials  show: 

WIDTH  IN  LENGTH 

M.  olivacea  M.  carolinensis 

Males  1.65-4.0  av.  2.16  I»7"3-1  av-  2-2 

Females  2.46-3.1  av.  2.7  1-83-2.3  av.  2.0 

The  female  M.  carolinensis  is  certainly  wider  in  body  than  M.  olivacea.  For 
example:  A  ripe  female  of  the  former  30  mm.  long  is  16  mm.  wide,  one  of  the 
latter  30  mm.  long  is  12  mm.  wide;  or  a  ripe  female  of  the  former  27  mm. 
long  is  14.5  mm.  wide,  whereas  a  similar  one  of  the  latter  is  n  mm.  But  ap- 
parently width  into  length  is  not  a  good  distinguishing  character  for  the 
males.  The  most  constant  character  that  we  have  found  is  Girard's  depressed 
form  of  the  body.  This  is  certainly  a  flattened  frog.  Our  measurements  are: 

THICKNESS  INTO  LENGTH 

M.  olivacea  M.  carolinensis 

Males:  3-0-5-6,  av.  3.7  2-5-3-35>  av-  2&5 

Females:  4-o-4-3>  av.  4.2  2.65-3.4,  av.  2.9 

(in  2  ripe  females,         (in  2  ripe  females, 
4.2, 4.0)  2.7,  2.65) 

In  the  field  in  March,  1925,  we  wrote  that  we  believed  they  had  more  poinfed 
heads  than  M .  carolinensis.  We  now  find  that  this  impression  is  due  to  the 
more  depressed  head. 
Limbs  are  more  slender  in  M.  olivacea. 

TIBIA  WIDTH  INTO  LENGTH 
M.  olivacea  M.  carolinensis 

3-4  21/2-3 

The  latter  has  thicker  tibia  and  femur. 


NARROW-MOUTH  TOADS:  BREVICIPITIDAE  581 

"Slate  gray  upper  parts,  usually  with  some  indefinitely  outlined  small  black 
spots  concentrated  on  middorsal  region  and  hind  quarters;  ventral  surfaces 
white,  immaculate,  occasionally  with  very  faint  reticulations  in  gular  region 
and  on  sides;  skin  smooth. 

".  .  .  From  carolinensis,  olivacea  differs  largely  in  color,  the  southeastern 
species  being  usually  brownish  above,  sometimes  reticulated  with  darker 
brown,  and  at  other  times  with  an  indistinct  broad  band  on  the  back  extending 
from  the  post-orbital  region  to  the  hind  legs,  about  as  broad  as  the  interorbital 
distance  anteriorly,  gradually  widening  posteriorly  until  at  the  groin  it  covers 
%  of  the  back.  This  particular  marking  is  very  suggestive  of  Gastrophryne 
elegans  of  Mexico  and  Central  America.  The  ventral  surfaces  of  carolinensis 
are  distinctly  reticulated  with  light  brown"  (H.  M.  Smith,  Kans.,  1934,  pp. 
501-502). 

"Engystoma  olivacettm  nob.  There  is  a  single  specimen  of  Engystoma,  one 
inch  one  line  in  length  by  seven  lines  in  breadth;  the  anterior  extremities  6; 
the  posterior  i  inch  10  lines  in  length.  The  general  color  above  is  olive,  with 
dark  colored  spots  posteriorly;  these  are  observed  in  considerable  number 
upon  the  posterior  part  of  the  body  and  upper  parts  of  the  thighs;  a  few  also 
are  seen  over  the  shoulders;  the  sides  are  obscurely  marbled  with  brown; 
under  surface  yellow,  immaculate;  a  well  marked  fold  passes  across  the  head 
behind  the  eyes,  extending  down  alongside  the  head. 

"This  specimen  is  larger  than  any  of  those  in  the  collection  of  the  Academy, 
and  the  coloration  is  quite  different  from  that  of  Dr.  Holbrook's  figure  .  .  . 
and  the  specimens  from  Georgia  in  the  collection  presented  by  Maj.  Le  Conte. 
These  are  all  more  or  less  mottled  with  brown  beneath  and  brown  and  chest- 
nut colored  above.  Dimensions:  As  above,  length  of  tarsus  and  toes  9  lines; 
of  leg  4  lines;  of  thigh  4;  of  arm  2,  or  forearm  2.  Structure:  II.  Engystoma 
texense. — Head  constituting  the  third  of  the  total  length.  Snout  forming  an 
acute  triangle  rounded  at  the  summit.  Gape  of  the  mouth  horizonal,  its  cleft 
extending  to  a  perpendicular  line  drawn  posteriorly  to  the  pupil.  Limbs 
slender;  three  carpal  callosities,  and  one  metatarsal  tubercle.  Palm  of  hands 
and  sole  of  feet  smooth.  Skin  smooth  also.  Color  above  light  olivaceous  Jsrown, 
with  a  few  black  dots  posteriorly;  beneath  uniform  whitish. 

"This  species  is  allied  to  B.  carolinense,  and  differs  from  it  by  a  more  de- 
pressed and  flattened  head,  a  more  truncated  snout,  which,  as  usual,  protrudes 
beyond  the  lower  jaw.  The  body  itself  is  likewise  more  depressed,  and  the 
limbs  assume  a  slender  appearance. 

"The  head  is  continuous  with  the  body,  and  constitutes  about  the  third  of 
their  combined  length.  The  cleft  of  the  mouth  does  not  extend  as  far  back  as 
in  E.  carolinense,  since  it  corresponds  to  a  perpendicular  line  drawn  behind 
the  pupil.  The  longitudinal  diameter  of  the  eye  is  equal  to  the  distance  be- 
tween the  orbit  and  the  nostril.  The  interocular  space  measured  across  the 
anterior  rim  of  the  orbits,  is  greater  than  the  rostral  space  from  the  orbits  for- 


582  HANDBOOK  OF  FROGS  AND  TOADS 

wards.  The  symphysis  of  the  lower  jaw  presents  the  same  structure  as  in  the 
species  just  alluded  to. 

"The  skin  is  perfectly  smooth  throughout  in  all  the  specimens  which  we 
have  examined;  they  were  collected  in  April,  and  are,  no  doubt,  liable  to  as- 
sume a  rougher  appearance  during  the  hotter  and  more  dry  season  of  the 
year. 

"Gen.  Remarks.  Dumeril  and  Bibron  describe  five  species  of  Engystoma — 
two  from  N.  America  (carohnense  and  rugosum),  two  from  S.  America 
(ovalc  and  micropi),  and  one  from  Malabar  (ornatum).  Habitat.  Accord- 
ing to  Dr.  Holbrook,  Engystoma  carohnense  has  never  been  found  north  of 
Charleston,  its  range  extending  westward  to  the  lower  Mississippi"  (E.  Hallo- 
well,  Gen.,  18563,  VIII,  252). 

"The  ground  color  of  the  upper  region  of  the  head,  body,  and  limbs,  is  of  a 
light  olivaceous  brown  tint,  anteriorly  uniform,  posteriorly  besprinkled  with 
small  black  spots  or  dots  especially  over  the  coccyx  and  thighs.  A  whitish  tint 
pervades  uniformly  throughout  the  inferior  regions.  Specimens  of  this  species 
were  procured  in  Texas,  by  Capt.  John  Pope"  (C.  Girard,  Gen.,  1859,  169- 
170). 

Voice:  This  frog  begins  its  call  with  a  pleasant  little  whistle  and  then  runs 
into  its  bleat.  It  opens  something  like  whee. 

When  we  first  heard  these  frogs  8:30-9:50  P.M.,  we  thought  their  call  dif- 
ferent from  that  of  our  eastern  ones,  a  much  lower  and  less  carrying  note  than 
that  of  M.  carolmensis. 

"The  breeding  call  is  a  high,  shrill  buzz  of  some  2-3  seconds  duration,  and 
of  such  slight  volume  that  a  single  call  cannot  be  heard  more  than  fifty  to  a 
hundred  feet  away.  A  full  chorus  sounds  like  a  band  saw  in  operation"  (H.  M. 
Smith,  Kans.,  1934,  p.  503).  "The  note  of  this  species  is  a  shrill  long-drawn 
'quaw,  quaw,5  repeated  at  intervals  of  several  seconds  duration.  This  note  is 
quite  different  from  the  loud  explosive  'quack*  of  Bufo  compactilis"  (J.  K. 
Strecker,  Jr.,  Tex.,  1926^  p.  12). 

Breeding:  This  species  breeds  from  March  15  to  September  in  heavy  rain 
periods.  The  egg  mass  is  a  surface  film,  the  complement  645,  the  eggs,  black 
and  white.  The  egg  is  %o-%r>  inch  (0.8-0.9  mm.);  the  envelope  %~/^  inch 
(2.8-3.0  mm.),  loose  and  irregular,  possibly  merging  in  the  film  mass.  The 
grayish  olive  tadpole  is  small,  1{Kc>  inch  (23  mm.),  flat  and  wide,  the  tail* tip 
black,  the  eye  just  visible  from  the  ventral  aspect.  There  are  no  teeth,  no 
horny  mandibles,  and  no  papillae.  After  30  to  50  days,  the  tadpoles  transform 
from  April  15  to  October  at  %-%  inch  (10-12  mm.).  Amplexation  as  in 
M.  carolinensts. 

April  15,  Beeville,  Tex.,  cattletank  area.  Microhyla  tadpoles  are  at  the  sur- 
face among  brush.  Some  are  mature,  some  with  tails  getting  short,  some  with 
hind  legs  well  developed.  They  ought  to  transform  in  a  week  or  two.  In  the 
dirty  water,  when  one  comes  to  rest  at  the  surface,  his  circular  body  and  black 


NARROW-MOUTH  TOADS:  BREVICIPITIDAE  583 

tail  tip  are  very  prominent.  They  seem  to  have  more  speckled  tails  than  M. 
carolinensis.  They  seem  to  be  more  or  less  localized  in  the  brushy  areas. 

"Little  seems  to  have  been  recorded  of  the  breeding  habits  of  this  frog. 
Dickerson  (1913,  p.  169)  remarks  that  they  were  found  breeding  during  late 
June  in  1905  near  Brownsville,  Texas.  In  early  September  of  1932  I  heard  large 
choruses  in  the  same  region.  The  season  in  Kansas  may  be  extremely  late,  as 
a  female  collected  during  August  of  1926  in  Montgomery  County  near  In- 
dependence retained  large  numbers  of  eggs  in  her  abdomen.  Other  females 
collected  as  late  as  June  16  near  Manhattan  contained  eggs"  (H.  M.  Smith, 
Kans.,  1934,^503). 

"Again,  hatching  in  Microhyla  olivacea  is  more  like  Hyla  v.  versicolor  than 
any  other  so  far  observed,  although  these  forms  belong  to  different  families" 
(A.  N.  Bragg,  Okla.,  1944,  p.  234). 

Journal  notes:  March  24, 1925,  Beeville,  Tex.  In  a  roadside  ditch,  Microhyla 
are  at  the  edge,  above  the  water,  croaking.  We  approached  one  slowly,  didn't 
see  him  croak,  but  the  instant  the  light  was  put  on  him,  he  began  to  crawl 
up  the  bank  and  through  the  grass,  mouselike,  going  very  fast. 

June  15,  1930,  Beeville,  Tex.  Heard  only  one  or  two.  Took  me  an  hour  to 
locate  one  of  three  croaking.  Much  less  volume  than  in  M.  carolinensis.  Not 
so  much  bleat  in  its  note.  They  must  be  over  breeding  in  this  region.  Can 
find  no  eggs  or  tadpoles  in  old  places  of  5  years  ago. 

July  10,  1925,  Fort  Davis  Mts.,  Tex.  In  a  large  permanent  pool  2  miles 
south  of  Fort  Davis,  we  were  much  surprised  to  find  one  lone  narrow- 
mouthed  toad  tadpole,  a  mature  one.  This  was  the  only  one  we  caught.  Strecker 
credits  the  Texas  narrow-mouth  to  east-central,  central,  and  southern  Texas. 
We  know  of  no  record  of  Microhyla  this  far  westward.  In  the  present  state 
of  our  knowledge  of  Microhyla  tadpoles,  it  looks  to  be  similar  to  M.  olivacea 
larvae  which  we  took  at  Beeville  and  San  Antonio.  This  seems  to  justify  Stej- 
neger  and  Barbour  in  giving  the  range  of  M.  olivacea  as  Texas. 

July  22,  1942,  Toyahvale,  Tex.  Night  trip  west  on  highway.  In  this  pond 
heard  one  lone  Microhyla  olivacea.  Farther  on  heard  plenty  of  Microhyla. 
Then  I  began  to  question  in  my  mind  whether  they  were  all  this  species.  (See 
Bufo  insidior,  p.  185.) 

Authorities'  corner: 
H.  W.  Parker,  Gen.,  1934,  pp.  1-208, 
W.  F.  Blair,  Okla.,  1936,  p.  115. 
G.  A.  Moore  and  C.  C.  Rigney,  Okla.,  1942,  p.  78, 


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1929  Patch,  C.  L.  Copeia,  no.  4,  p.  235. 

1929  Barbour,  T.,  and  A.  Loveridge.  Bull.  Mus.  Comp.  Zool.f  69™,  205-360. 

1929  Wright,  A.  H.  Proc.  U.S.  Nat.  Mus.,  74,  art.  XI,  1-70. 

1930  Metcalf,  Z.  P.  A.  Textboo^  of  Economic  Zoology.  Philadelphia.  Pp.  1-392. 

1930  Netting,  M.  G.  Papers  Mich.  Acad.  Sci.  Arts  Letters  (1929),  pp.  437-443. 

1931  Ahl,  E.  In  Amphibia  (Lurche)  Reptilia  (Kriechtiere)  Tabulae  Biologicae, 
ed.  W.  Junk.  Berlin.  Suppl.  2  (1930),  pp.  598-715;  Tab.  biol.  period  i 

(i930>PP-  346-384-  f          f       ^ 

1931    Noble,  G.  K.  The  Biology  of  the  Amphibia.  New  York  and  London.  Pp. 


- 
1932    Wright,  A.  H.  Life  Histories  of  the  Frogs  of  Ofefinofee  Swamp,  Georgia. 

New  York.  Pp.  1-497. 

1932  Kellogg,  R.  Bull.  U.S.  Nat.  Mus.,  no.  160,  pp.  1-224. 
19333    Burt,  C.  E.  Amer.  Midi.  Naturalist,  14*,  35°-354- 

1933  Villicrs,  C.  G.  S.  de.  Nature,  i3i3315,  693-694. 

1934  Villicrs,  C.  G.  S.  de.  Studies  .  .  .  Ascaphus  truei.  .  .  .  Bull.  Amer.  Mus. 
Nat.  Hist.,  77,  1-18. 

1934  Parker,  H.  W.  A  Monograph  of  the  Frogs  of  the  Family  Microhylidae. 
British  Museum,  London.  Pp.  1-208. 

1935  Burt,  C.  E.  Amer.  Midi.  Naturalist,  i63,  31  1-319. 
1935     Harper,  F.  Same,  i6J,  275-310. 

1935    Lm,  C.  C.  Jour.  Morphol.,  57,  131-142.  (See  Schmidt,  K.  P.,  Copeia,  no.  4, 
1938,  p.  99.) 

1935  Wright,  A.  H.  Proc.  Nat.  Acad.  Sci.  (Washington),  21°,  34°"345- 

1936  Bragg,  A.  N.  Amer.  Naturalist,  70,  459-466. 

1936  Kauffeld,  C.  F.  Herpetologica,  i1,  1  1. 

19363    Taylor,  E.  H.  Kans.  Univ.  Sci.  Bull.,  2420,  505-516.  (Bull.  Univ.  of  Kans.f 
3714,  issued  Feb.  16,  1938.) 

1937  Gaige>  H-  T-»  N-  Hartweg,  and  L.  C.  Stuart.  Occ.  Papers  Mus.  Zool.  Univ. 
Mich.,  no.  357,  pp.  1-6. 

1937    Gaige,  H.  T.  Univ.  Mich.  Studies  Sci.  Ser.,  12,  301-302  (Tamauhpas). 

1937    Kauffeld,  C.  F.  Herpetologica,  i3,  84-86. 

1937    Oliver,  J.  A.  Occ.  Papers  Mus.  Zool.  Univ.  Mich.,  no.  360,  pp.  1-8. 


588  HANDBOOK  OF  FROGS  AND  TOADS 

1938    Browncll,  L.  W.  Amer.  Photography,  32,  353-354. 

1938    Burt,  C.  E.  Trans.  Kans.  Acad.  Sci.,  41, 331-360. 

1938    Gloyd,  H.  K.  Turtox  News,  16%  49-53,  66-67. 

1938    Schmidt,  K.  P.  Copeia,  no.  4,  p.  199. 

i938a    Schmidt,  K.  P.  Field  Mus.  Nat.  Hist.,  Zool.  ser.,  2029, 377-382. 

1938  Taylor,  E.  H.  Bull.  Univ.  of  Kans.,  3911.  (Kans.  Univ.  Sci.  Bull,  25™,  385- 
405.) 

1939  Bushnell,  R.  J.,  E.  P.  Bushnell,  and  M.  V.  Parker.  Jour.  Tenn.  Acad.  Sci.,  14*, 
209-215.  (Chromosome  study.) 

1939  DcSola,  R.  Reptiles  and  Amphibians,  an  Illustrated  Natural  History.  (Fed- 
eral Writers'  Project,  W.P.A.,  New  York  City.)  Chicago. 

1939  Dunn,  E.  R.  Proc.  Acad.  Nat.  Sci.  Phila.,  90,  153-154. 

1939  Eaton,  T.  H.,  Jr.  Copeia,  no.  2,  pp.  95-97. 

1939  Janes,  R.  G.  Copeia,  no.  3,  pp.  134-140. 

1939  McClung,  C.  E.  Tabulae  Biologicac,  ed.  W.  Junk.  Vol.  XVIII,  pt.  1, 39-47. 

1939  Morgan,  A.  H.  Animals  in  Winter.  New  York.  Pp.  331-369. 

1939  Tanner,  V.  M.  Great  Basin  Naturalist,  i1,  3-26. 

1940  Babcock,  H.  L.,  and  I.  Hoopes.  New  Eng.  Naturalist,  no.  6,  pp.  7-9. 
1940  Cochran,  D.  M.  Nature  Magazine,  33, 324. 

1940  Cott,  H.  B.  Adaptive  Coloration  in  Animals.  London.  Pp.  1-508. 

1940  Devoe,  A.  Amer.  Mercury,  5i199,  365-368. 

1940  Dunn,  E.  R.,  and  M.  T.  Dunn.  Copeia,  no.  2,  pp.  69-74  (esp.  pp.  71-72). 

1940  Harper,  F.  Amer.  Midi.  Naturalist,  23%  692-723. 

1940  Moore,  J.  A.  Amer.  Naturalist,  74,  89-93. 

1940  Necker,  W.  L.  Chicago  Acad.  Sci.,  Leaflet  no.  15,  pp.  1-2. 

1940  Necker,  W.  L.  Same,  no.  16,  pp.  i,  2. 

1940  Rugh,  R.,  and  I.  Exner.  Proc.  Amer.  Phil.  Soc.t  no.  5,  pp.  607-619. 

1940  Smith,  A.  L.  Jour.  Biol.  Photeg.  Assoc.,  9%  11-17. 

1940  Stejneger,  L.  Copeia,  no.  3,  pp.  149-151. 

1941  lllingworth,  J,  T.  Proc.  Ent.  Soc.  (1940),  n,  51. 
1941  Moore,  J.  A.  Jour.  Exp.  Zool.,  863, 405-422. 

1941     Raney,  E.  C.,  and  W.  M.  Ingram.  Amer.  Midi.  Naturalist,  26*,  201-206. 
1941     Rugh,  R.  Proc.  Amer.  Phil.  Soc.,  34°,  617-631. 

1941  Smith,  C.  C.,  and  A.  N.  Bragg.  Anat.  Rec.,  8i4,  suppl.  p.  in. 
i94ia    Smith,  C.  C.,  and  A.  N.  Bragg.  Same,  p.  53. 

i942g    Bragg,  A.  N.  Anat.  Rec.,  84^  507. 

1942  Colliers,  Feb.  7,  1942,  pp.  60-61. 

1942    DeVoe,  A.  Amer.  Mercury,  54220,  491-495. 

1942  Driver,  E.  C.  Name  That  Animal  .  .  .  Amphibians  and  Reptiles.  North- 
ampton, Mass.  Pp.  293-413. 

19423    Moore,  J.  A.  Biological  Symposia.  Lancaster,  Pa.  Vol.  VI,  pp.  189-213. 

1942    Coin,  C.  J.  Copeia,  no.  3,  pp.  183-184. 

1942    Mittleman,  M.  B.,  and  H.  T.  Gier.  Proc.  N.  E.  Zool.  Club,  20, 7-15. 

1942    Moore,  J.  A.  Biol.  Bull.,  no.  83,  pp.  375-388. 

1942  Noble,  G.  K.,  and  L.  R.  Aronson.  Bull.  Amer.  Mus.  Nat.  Hist.,  80,  art.  V, 
127-142. 


BIBLIOGRAPHY 


589 


1942  Schmidt,  K.  P.  Copeia,  no.  i,  pp.  53-54. 

19433  Blair,  A.  P.  Amer.  Midi.  Naturalist,  pp.  615-620. 

1943  Lynn,  W.  G.,  and  J.  N.  Dent.  Copeia,  no.  4,  pp.  234-242. 
1943  Preble,  E.  A.  Nature  Magazine,  36*',  137-139. 

1943  Pusey,  H.  K.  Quart.  Jour.  Micros.  Sci.,  new  ser.,  S^l~BA\  105-185. 

1943  Taylor,  E.  H.  Bull.  Univ.  of  Kans.,  29%  pt.  2,  343-361. 

1944  Lynn,  W.  G.  Copeia,  no.  3,  pp.  189, 190. 

1944  Marr,  J.  C.  Amer.  Midi.  Naturalist,  yp,  478-481. 

1944  Moore,  J.  A.  Bull.  Amer.  Mus.  Nat.  Hist.,  82%  345-370. 

1944  Palmer,  E.  L.  Nature  Magazine,  37°,  305-312. 

1945  Bogert,  C.  M.,  and  J.  A.  Oliver.  Bull.  Amer.  Mus.  Nat.  Hist.,  83,  art.  6,  301- 
425. 

1945  Brownell,  L.  W.  Amer.  Photography,  39%  34-37. 

1945  Ely,  C.  A.  Copeia,  no.  4,  p.  256. 

1945  Livezey,  R.  L.,  and  A.  H.  Wright.  Amer.  Midi.  Naturalist,  34%  701-706. 

1945  Mittleman,  M.  B.  Copeia,  no.  i,  pp.  31-37. 

1945  Smith,  H.  M.  Ann.  Carnegie  Mus.,  30,  89-92. 

1945  Taylor,  E.  H.,  and  H.  M.  Smith.  Proc.  U.S.  Nat.  Mus.,  95,  521-623. 

1946  Blair,  A.  P.  Amer.  Mus.  Novitates,  no.  1327,  pp.  1-3. 
1946  Bragg,  A.  N.  Herpetologica,  3,  pt.  $,  89-97. 

1946  Hecht,  M.  K.,  and  B.  L.  Matalas.  Amer.  Mus.  Novitates,  no.  1315,  pp.  1-21. 

1946  Mittleman,  M.  B.  Herpetologica,  3,  pt.  2,  57-60. 

1946  Netting,  M.  G.,  and  C.  J.  Gom.  Copeia,  no.  2,  p.  107. 

i947a  Blair,  A.  P.  Amer.  Mus.  Novitates,  no.  1344,  April  15,  1947. 

1947!)  Blair,  A.  P.  Copeia,  1947,  p.  67. 

1947  Harper,  F.  Proc.  Biol.  Soc.  Wash.,  60,  59-40. 

1947  Livezey,  R.  L.,  and  A.  H.  Wright.  Amer.  Midi.  Naturalist,  37%  177-222. 

1947  Richmond,  N.  D.  (see  Virginia). 

1947  Smith,  H.  M.  Herpetologica,  4, 7-13. 

II.  GENERAL  CHECK  LISTS  OR  CATALOGUES 

i875b    Cope,  E.  D.  Bull.  U.S.  Nat.  Mus.,  no.  i,  pp.  1-10,  29-32,  55-104. 

1882  Yarrow,  H.  C.  Same,  no.  24,  pp.  1-249. 

1883  Davis,  N.  S.,  Jr.,  and  F.  L.  Rice.  Bull.  111.  State  Lab.  Nat.  Hist.,  no.  5,  pp. 
15-25. 

1884  Carman,  S.  Bull.  Essex  Inst.,  i6l~3,  41-46. 
1886    Cope,  E.  D.  Proc.  Amer.  Phil.  Soc.,  23, 514-521. 

1917-1943  Stejneger,  L.,  and  T.  Barbour.  A  Chec\  List  of  North  American  Am- 
phibians and  Reptiles.  Cambridge,  ist  ed.,  1917.  2d  ed.,  1923.  3d  ed.,  1933. 
4th  ed.,  1939.  5th  ed.,  1943. 

1945    Taylor,  E.  H.,  and  H.  M.  Smith.  Proc.  U.S.  Nat.  Mus.,  95,  521-623. 

1947  Logier,  E.  B.  S.,  and  G.  C.  Toner.  Chec\  List  of  Canadian  Amphibians  and 
Reptiles.  Manuscript. 

1948  Smith,  H.  M.,  and  E.  H.  Taylor.  An  Annotated  Checklist  and  Key  to  the 
Amphibia  of  Mexico.  Bull.  U.S.  Nat.  Mus.,  no.  194,  pp.  1-118. 


590  HANDBOOK  OF  FROGS  AND  TOADS 

III.  STATE  AND  PROVINCE  LISTS 
CANADA 

Alberta 

1908    Preble,  E.  A.  North  Amer.  Fauna,  no.  27,  pp.  501-502.  Washington,  D.C 

1929    Patch,  C.  L.  Can.  Field  Naturalist,  43, 137-138. 

1931    Harper,  F.  Same,  45,  68-70. 

1931    Logier,  E.  B.  S.  Same,  45,  90. 

1939    Patch,  C.  L.  (see  Ontario). 

1942    Logier,  E.  B.  S.,  and  G.  C.  Toner.  Can.  Field  Naturalist,  56,  15-16. 

1942  Slater,  J.  R.  (see  Washington). 

1943  Slipp,  J.  W.,  and  G.  C.  Carl  (sec  British  Columbia). 
1946    Williams,  M.  Y.  Can.  Field  Naturalist,  6, 47-49. 

Athabaska 

1908    Preble,  E.  A.  North  American  Fauna,  no.  27,  pp.  501-502. 

1931     Harper,  F.  (see  Alberta). 
British  Columbia 

1894    Cope,  E.  D.  Proc.  Acad.  Nat.  Sci.  Phila.  (1893),  45,  181-184. 

1898    Fannin,  J.  A  Preliminary  Catalogue  of  the  Collections  of  Natural  His 
tory  and  Ethnology  in  the  Provincial  Museum.  Victoria,  B.C.  P.  58. 

1922    Patch,  C.  L.  Copeia,  1922,  pp.  76-78. 

1926  Hardy,  G.  A.  Rept.  Prov.  Mus.  Nat.  Hist.  B.C.  (1925),  pp.  C23-C24. 

1927  Hardy,  G.  A.  Same  (1926),  pp.  C25,  C$g. 

1928  Hardy,  G.  A.  Same  (1927),  pp.  £17,  19. 

1928  Slevm,  J.  R.  Occ.  Papers  Calif.  Acad.  Sciences,  16,  1-152. 

1929  Newcome,  W.  A.  Rept.  Prov.  Mus.  Nat.  Hist.  B.C.  ( 1928),  p.  F24. 

1929  Patch,  C.  L.  (see  Alberta). 

1930  Newcome,  W.  A.  Rept.  Prov.  Mus.  Nat.  Hist.  B.C.  (1929),  p.  F22. 

1931  Newcome,  W.  A.  Same  (1930),  p.  Gig. 

1932  Logier,  E.  B.  S.  Trans.  Roy.  Can.  Inst.,  18,  pt.  2,  311-315,  319-324. 

1933  Williams,  M.  Y.  Rept.  Prov.  Mus.  Nat.  Hist.  B.C.  ( 1932),  p.  C22. 
1935     Ricker,  W.  E.,  and  E.  B.  S.  Logier.  Copeia,  no.  i,  p.  46. 

1937    Cowan,  L  McT.  Rept.  Prov.  Mus.  Nat.  Hist.  B.C.  (1936),  pp.  16-25. 

1937    Cowan,  I.  McT.  Same  (1936),  pp.  Ki8-2o. 

1939    Cowan,  I.  McT.  Occ.  Papers  Brit.  Col.  Prov.  Mus.,  i,  92-93. 

1939    Patch,  C.  L.  (see  Ontario). 

1941  Cowan  I.  McT.  Copeia,  no.  i,  p.  48. 

1942  Carl,  G.  C.  Copeia,  no.  2,  p.  129. 

1942  Slater,  J.  R.  (see  Washington). 

1943  Carl,  G.  C.  Brit.  Col.  Prov.  Mus.  Handboo\,  no.  2,  pp.  1-62. 
1943    Slipp,  J.  W.,  and  G.  C.  Carl.  Copeia,  no.  2,  p.  127. 

1945    Carl,  G.  C.,  and  I.  McT.  Cowan.  Copeia,  no.  i,  p.  43. 

19453    Carl,  G.  C.,  and  I.  McT.  Cowan.  Copeia,  no.  i,  pp.  52-53. 
Keewatin 

1902    Preble,  E.  A.  North  Amer.  Fauna,  no.  22,  pp.  133-134- 
Labrador 

1925    Speck,  F.  G.  Copeia,  no.  138,  pp.  5-6. 


BIBLIOGRAPHY 


591 


1938    Schmidt,  K.  P.  (see  Illinois). 
Mackenzie 

1908    Preble,  E.  A.  (see  Alberta). 

1931     Harper,  F.  (see  Alberta), 
Manitoba 

1918    Seton,  E.  T.  Ottawa  Naturalist,  32,  82-83. 

1921  O'Donoghue,  C.  H.  Can.  Field  Naturalist,  35,  121-128,  129-131. 

1938  Schmidt,  K.  P.  (see  Illinois). 

1939  Patch,  C,  L.  (see  Ontario). 
Newfoundland 

1926    Johansen,  Frits.  Can.  Field  Naturalist,  40, 16. 
New  Brunswick 

1873    Adams,  A.  L.  Field  and  Forest  Rambles.  London.  Pp.  197,  303. 

1898  Cox,  P.  Nat.  Hist.  Soc.  New  Bruns.t  4,  pt.  i,  art.  VI,  64-66. 
18993    Cox,  P.  Trans.  Roy.  Soc.  Can.,  2d  ser.  5,  sec.  IV,  art.  V,  152,  153. 
iSggb    Cox,  P.  Ottawa  Naturalist,  13,  194,  195. 

1939    Patch,  C.  L.  (see  Ontario). 
Nova  Scotia 

1865    Jones,  J.  M.  Proc.  and  Trans.  Nova  Scotia  Inst.  Sci.,  i  (not  2),  pt.  Ill 
(1864-1865).  Halifax.  Pp.  116,  122-126. 

1896    Mackay,  A.  H.  Proc.  and  Trans.  Nova  Scotia  Inst.  Sci.,  9  (2d  ser.,  2),  xli- 
xliii. 

18993    Cox,  P.  (see  New  Brunswic^). 

iSggb    Cox,  P.  (see  New  BrunswicJ(). 

1939    Patch,  C.  L.  (see  Ontario). 
Ontario 

1850    Agassiz,  A.  LaJ(e  Superior.  Boston.  Pp.  378-380,  382. 

1858    Agassiz,  A.  In  The  Handboo\  of  Toronto.  Toronto,  1878.  Pp.  60-61. 

1878    Yarrow,  H.  C.,  and  H.  W.  Henshaw  (see  Nevada). 

1881     Gamier,  J.  H.  The  Canadian  Sportsman  and  Naturalist,  i5,  38,  39. 

1883  Gamier,  J.  H.  Amer.  Naturalist,  17°,  945-954. 

1884  Small,  H.  B.,  and  William  P.  Lett.  Ottawa  Field-Naturalist's  Club  Trans., 
No.  5,  21,  148,  149. 

1885  Small,  H.  B.,  and  William  P.  Lett.  Same,  No.  6,  211,  282-283. 

1899  Meek,  S.  E.  Field  Columbian  Mus.,  Pub.  no.  41,  i17,  331. 

1902    Meek,  S.  E.,  and  H.  W.  Clark.  Same,  no.  67,  Zool.  ser.,  3%  139-140- 
1908    Nash,  C.  W.  In  Vertebrates  of  Ontario  (Dept.  of  Education,  Toronto). 

"Batrachians  and  Reptiles  of  Ontario,"  pp.  7-10. 
1913    Piersol,  M.  H.  In  J.  H.  Faull,  Natural  History  of  the  Toronto  Region. 

Toronto.  Ch.  xviii,  "Amphibia,"  pp.  244-248. 
1915    Wright,  A.  H.  Copeia,  no.  23,  pp.  46-48. 
1918    Patch,  C.  L.  Ottawa  Naturalist,  32*,  53. 
1920    Williams,  M.  Y.  Can.  Field  Naturalist,  34%  125. 
1920    Wright,  A.  H.,  and  S.  E.  R.  Simpson.  Same,  34,  144. 

1922  Klugh,  A.  B.  Copeia,  no.  103,  pp.  14,  15. 

1925    Logier,  E.  B.  S.  Can.  Field  Naturalist,  39%  91-93. 
1928    Brown,  J.  R.  Can.  Field  Naturalist,  42, 126. 


592  HANDBOOK  OF  FROGS  AND  TOADS 

1928    Dymond,  J.  R.  Cont.  Roy.  Ont.  Mus.  ZooL,  no.  2  (reprinted  from  Univ. 

Toronto,  Studies  Biol.,  ser.  32),  pp.  35-36. 
1928    Logier,  E.  B.  S.  Trans.  Roy.  Can.  Inst.,  i62,  279-291. 
1931     Coventry,  A.  F.  Can.  Field  Naturalist,  45,  109. 

1931  Logier,  E.  B.  S.  Trans.  Roy.  Can.  Inst.,  18,  pt.  i,  no.  39,  229-232. 

1932  Gaige,  H.  T.  Copeia,  no.  3,  p.  134. 

1934    Logier,  E.  B.  S.  In  R.  D.  Black's  "Chas.  Fothergill's  Notes  .  .  .  ,"  Trans. 
Roy.  Can.  Inst.,  2O43,  pt.  I,  160. 

1937  Logier,  E.  B.  S.  Handb.  3,  Roy.  Ont.  Mus.  ZooL,  pp.  1-16  (see  School, 
Apr.,  May,  June,  1937). 

1938  Toner,  G.  C.,  and  W.  E.  Edwards.  Can.  Field  Naturalist,  52,  40-43  (esp. 
42). 

1939  Patch,  C.  L.  Copeia,  no.  4,  p.  235. 

1940  Allin,  A.  E.  Trans.  Roy.  Can.  Inst.,  23,  83-1 1 8. 

1941  Logier,  E.  B.  S.,  Univ.  Toronto,  Studies  Biol.  Scr.,  48,  pp.  93-101. 

1941  Toner,  G.  C.,  and  N.  de  St.  Remy.  Copeia,  no.  i,  pp.  10-13. 

1942  Logier,  E.  B.  S.  Cont.  Roy.  Ont.  Mus.  ZooL,  no.  21,  pp.  154-163. 

1942  Williams,  M.  Y.  Can.  Field  Naturalist,  56,  no.  6,  p.  93. 

1943  Logier,  E.  B.  S.,  and  G.  C.  Toner.  Can.  Field  Naturalist,  57,  104-105. 
Prince  Edward  Island 

18993    Cox,  P.  (sec  New  Brunswic^). 
i899b    Cox,  P.  (see  New  Brunswick). 
Quebec 

1874-1875    Provancher,  L.  Nat.  Can.,  6,  273-278;  7,  10-20,  42-46,  65-73,  2%9~~ 

298*  32I~33<>>  353-37°- 
iSQga    Cox,  P.  (see  New  Brunswick). 

1899!)    Cox,  P.  (see  New  Brunswic^). 

1937  Ball,  S.  C.  Copeia,  no.  4,  p.  230. 

1938  Trapido,  H.,  and  R.  T.  Clausen.  Copeia,  no.  $,  pp.  117-125. 

1939  Moore,  J.,  and  B.  Moore.  Copeia,  no.  2,  p.  104. 
1939    Patch,  C.  L.  (see  Ontario). 

1941  Grant,  R.,  Copeia,  no.  3,  pp.  151-153. 

1941  Vladykov,  V.  D.,  Can.  Field  Naturalist,  55°,  pp.  83-84. 
Saskatchewan 

1908  Preble,  E.  A.  (sec  Alberta). 

1931  Harper,  F.  (see  Alberta). 

1938  Schmidt,  K.  P.  (see  Illinois). 

1939  Patch,  C.  L.  (see  Ontario). 
1946  Williams,  M.  Y.  (see  Alberta). 

THE  UNITED  STATES 

Alabama 

1910    Brimley,  C.  S.  Proc.  Biol.  Soc.  Wash.,  23,  9-18. 
1920    Dunn,  E.  R.  Same,  33, 129-130, 135-136. 

1922    Loding,  H.  P.  Geol.  Survey,  Alabama  (Mus.  Paper  no.  5,  Ala.  Mus.  Nat. 
Hist.  Univ.  Ala.),  pp.  16-21. 


BIBLIOGRAPHY  593 

1924  Holt,  E.  G.  Copeia,  no.  135,  p.  95. 

1931  Haltom,  W.  L.  Mus.  Papers,  Ala.  Mus.  Nat.  Hist.,  u,  1-145. 

1938  Viosca,  P.  Copeia,  no.  4,  p.  201. 

1939  Cahn,  A.  R.  (see  Tennessee). 

1940  Penn,  G.  H.,  Jr.  Jour.  Tenn.  Acad.  Sci.t  15%  353-355. 
1942  Netting,  M.  G.,  and  C.  J.  Coin.  Copeia,  no.  4,  p.  259. 

Alaska 

1898  Van  Dcnburgh,  J.  Proc.  Amer.  PAH.  Soc.,  37, 139. 

1920  Dice,  L.  R.  Occ.  Papers  Mus.  Zool.  Univ.  Mich.,  no.  85,  pp.  7, 1 1. 

1925  Williams,  M.  Y.  Can.  Field  Naturalist,  39%  72. 
1928  Slevin,  J.  R.  (see  British  Columbia). 

Arizona 

1859    Baird,  S.  F.  (see  Texas). 

1866    Cope,  E.  D.  Proc.  Acad.  Nat.  Sci.  Phtla.,  18,  301,  310-314. 

1875    Coues,  E.  In  G.  M.  Wheeler's  Kept.  Geog.  Explor.  West  oj  woth  Meridian. 

Washington.  Vol.  V,  "Zoology,"  627-631. 
1875    Yarrow,  H.  C.  In  same,  pp.  520-529. 

1890    Stejneger,  L.  North  American  Fauna,  no.  3,  art.  5,  pp.  116-118. 
1907    Mearns,  E.  A.  Bull.  U.S.  Nat.  Mus.,  no.  56,  pt,  i,  pp.  no,  113,  117,  118, 

125. 

1907    Ruthven,  A.  G.  Bull.  Amer.  Mus.  Nat.  Hist.,  23,  art.  XXIII,  483-51  r. 
1913    Van  Denburgh,  J.,  and  J.  R.  Slevin,  Proc.  Caltf.  Acad.  Set.,  4th  sen,  3, 

391-396. 
1917    Engelhardt,  G.  P.  Copeia,  no.  39,  pp.  5-7. 

1924  Cowles,  R.  A.  Jour.  Ent.  &  Zool.  Pomona  Coll.,  i64,  107-110. 

1925  Engelhardt,  G.  P.  Copeia,  no.  149,  pp.  91-92. 

1925  Ortenburger,  A.  I.  Univ.  of  Otya.  Bull.,  new  ser.,  no.  322,  Univ.  Studies 
no.  21.  (Proc.  Ofya.  Acad.  Set.,  4  [1924 1,  art.  VI,  19-20.) 

1926-1927  Ortenburger,  A.  I.,  and  R.  D.  Ortenburger.  Univ.  Otya.  Bull.,  new 
ser.,  no.  348,  Jan.  i,  1927.  (Proc.  Otya.  Acad.  Set.,  6  [1926],  pt.  1, 101-103.) 

1928    Slevin,  J.  R.  (see  British  Columbia). 

i92gb    Burt,  C.  E.,  and  M.  D.  Burt.  Jour.  Wash.  Acad.  Set.,  19,  428-434. 

1930  Musgrove,  M.  E.,  and  D.  M.  Cochran.  Copeia,  no.  173  (Oct.-Dec.  1929), 
Jan.  1 6,  1930,  pp.  97-99. 

1931  Campbell,  B.  Copeta,  no.  4,  p.  164. 

1931  Slevin,  J.  R.  Copeia,  no.  3,  pp.  140,  141. 

1932  Hobbs,  K.  L.  Copeia,  no.  2,  p.  104. 

i932a    Kellogg,  R.  Bull.  U.S.  Nat.  Mus.,  no.  160,  pp.  1-224. 

1932    King,  F.  W.  Copeta,  no.  2,  p.  99. 

1932    King,  F.  W.  Copeia,  no.  4,  pp.  175-176. 

1932    Klauber,  L.  M.  (see  California). 

i932a    Klauber,  L.  M.  Zool.  Soc.  San  Diego,  Bull.  9,  p.  77. 

1932  MacCoy,  C.  U.  Occ.  Papers  Boston  Soc.  Nat.  Hist.,  8, 11-15. 

1933  Allen,  M.  J.  Occ.  Papers  Mus.  Zool.  Univ.  Mich.f  no.  259,  pp.  1-3. 
19333    Burt,  C.  E.  (see  General  Worths). 

1933  Campbell,  B.  Copeia,  no.  2,  p.  100. 

1934  Campbell,  B.  Occ.  Papers  Mus.  Zool.  Univ.  Mich.,  no.  280,  pp.  1-9. 


594  HANDBOOK  OF  FROGS  AND  TOADS 

1934  McKee,  E.  D.,  and  C.  M.  Bogert.  Copeia,  no.  4,  p.  178. 
19353    Eaton,  T.  H.,  Jr.  Copeia,  no.  3,  p.  150. 

i935b    Eaton,  T.  H.,  Jr.  Rainbow  Bridge  Monument  Exped.  (1933)9  Bull.  3 
(June,  1935),  pp.  3-8. 

1935  Quaintance,  C.  W.  Copeia t  no.  4,  pp.  183,  184. 

1935  Wood,  W.R  (see  Utah). 

1936  Cowles,  R.  B.,  and  C.  M.  Bogert  (see  Nevada). 

1937  Dunn,  E.  R.  Proc.  Acad.  Nat.  Sci.  Phila.  (1936),  88,  471-472. 

1937  Gloyd,  H.  K.  Bull.  Chicago  Acad.  Sci.,  5,  79-136. 

1938  Dodge,  N.  N.  Grand  Canyon  Nat.  Hist.  Assoc.,  Nat.  Hist.  Bull.,  no.  9, 
pp.  1-15. 

1938  Taylor,  E.  H.  Bull.  Univ.  of  Kan.,  39'  %  421-445. 

1939  Chapel,  W.  L.  Copeia,  no.  4,  pp.  225-227. 

19403    Gloyd,  H.  K.  Chicago  Naturalist,  33~4,  67-78,  111-124  (esP-  i"-"3)- 

1940  Little,  E.  L.,  Jr.  Copeia,  no.  4,  pp.  260-265. 
1940    Tanner,  W.  W.  (see  Utah). 

1940    Taylor,  E.  H.,  and  I.  W.  Knobloch.  Proc.  Biol.  Soc.  Wash.,  53  (Oct.  7, 

1941),  125. 

1943    Arnold,  L.  W.  Copeia,  no.  2,  p.  128. 
1943    Kauffeld,  C.  K.  Amer.  Midi.  Naturalist,  29%  342-359. 
Arkansas 

1859    Baird,  S.  F.  Explor.  and  Surveys  R.  R.  Route  from  Miss.  River  to  Pacific 

Ocean  1853-56,  10,  Zool.  Rept.  no.  4,  37-45. 
1890  Cope,  E.  D.  Amer.  Naturalist,  24, 1204,  1205. 
1903  Stone,  W.  Proc.  Acad.  Nat.  Sci.  Phila.f  55,  538,  539. 

1908  Strecker,  J.  K.,  Jr.  Proc.  Biol.  Soc.  Wash.,  21,  88. 

1909  Hurter,  J.,  and  J.  K.  Strecker,  Jr.  Trans.  Acad.  Sci.  St.  Louis,  18,  11-15, 
18-20. 

1924  Strecker,  J.  K.,  Jr.  Baylor  Univ.  Bull.t  27%  pt.  3,  29-34. 

19293  Burt,  C.  E.,  and  M.  D.  Burt.  Amer.  Mus.  Novitates,  no.  381,  pp.  1-7. 

19293  Ortenburger,  A.  I.  (see  Oklahoma). 

1933  Black,  J.  D.  Copeia,  no.  2,  pp.  100,  101. 

1934  Perkins,  R.  M.,  and  M.  J.  R.  Lentz.  Copeia,  no.  3,  pp.  140,  189. 

1935  Burt,  C.  E.  (see  General  Worths). 

1935    Taylor,  E.  H.  Kans.  Univ.  Sci.  Bull.,  22,  207-218. 
1938    Black,  J.  D.  Copeia,  no.  i,  pp.  48, 49. 
California 

1853  Baird,  S.  F.,  and  C.  Girard.  Proc.  Acad.  Nat.  Sci.  Phila.,  6,  301,  302. 

1854  Baird,  S.  F.,  and  C.  Girard  (see  Oregon). 

1854    Hallowell,  E.,  Proc.  Acad.  Nat.  Sci.  Phila.,  6,  238. 

1856    Hallowell,  E.  Same,  7  (1854,  1855),  96,  97. 

1859    Baird,  S.  F.  (see  Texas). 

1859    Borland,  J.  N.  Proc.  Boston  Soc.  Nat.  Hist.  (1856-1859),  6,  193.  Read 
May  6, 1857. 

1859    Hallowell,  E.  Explor.  and  Surveys  Pacific  R.  R.  Route  1853-56.  Wash- 
ington, D.C.  Vol.  X,  Sen.  Ex.  Doc.  no.  78,  33d  Congress,  2d  session,  no.  i, 
1-23. 


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595 


1868  Cronise,  T.  F.  The  Natural  Wealth  of  California.  San  Francisco.  Pp. 
485,  486. 

1869  Cooper,  J.  G.  Proc.  Calif.  Acad.  Sci.,  4  (1868-1872).  San  Francisco,  1873. 
Pp.  61-81.  Read  Sept.  6,  1869. 

1869    Cooper,  J.  G.  Amei.  Natwahst,  3%  478,  480. 
1875    Yarrow,  H.  C.  (see  Arizona). 

1878  Yarrow,  H.  C.,  and  H.  W.  Henshaw  (see  Nevada). 

1879  Lockington,  W.  N.  Amer.  Naturalist,  13 !-,  780. 

1883  Stearns,  R.  E.  C.  Same,  17%  982. 

1884  Cope,  E.  D.  Proc.  Acad.  Nat.  Sci.  Phila.  (1883),  35,  15,  25-27,  32-35. 
1888    Cope,  E.  D.  Proc.  U.S.  Nat.  Mus.  (1887),  10,  241. 

1893     Stejneger,  L.  North  Amer.  Fauna,  no.  7,  pp.  159-161,  219-228. 
1907    Mearns,  E.  A.  Bull.  U.S.  Nat.  Mus.,  no.  56,  pt.  I,  pp.  133,  138. 
1912    Hurter,  }.  First  Ann.  Kept.  Laguna  Manne  Lab.  Claremont.  P.  67. 
I9i2a    Van  Denburgh,  J.  Proc.  Calif.  Acad.  Sa.t  4th  sen,  3  (Jan.  17,  1912),  149. 
1916    Camp,  C.  L.  Univ.  Calif.  Pub.  Zool.,  i21T,  503-512. 
I9i6a     Camp,  C.  L.  Same,  17°,  59-62. 

1916  Ruthlmg,  P.  D.  R.  Lorqutnia  (Los  Angeles),  i1, 6. 
J9i7b     Camp,  C.  L.  Univ.  Calif.  Pub.  ZooL,  17°,  115-125. 

1917  Gnnnell,  J.,  and  C.  L.  Camp.  Same,  17 lo,  127-208. 

1919    Hall,  H.  M.,  and  J.  Grinnell.  Proc.  Calif.  Acad.  Sci.,  4th  ser.,  9%  47,  54. 

1922  Van  Winkle,  K.  (see  Washington). 

1923  Marimon,  S.  Jour.  Ent.  &  Zool.,  15-,  27-31. 

1923  Storer,  T.  I.  Copcia,  no.  114,  p.  8. 

1924  Dunn,  E.  R.  Copcia,  no.  133,  pp.  75,  76. 

1924  Grinnell,  J.,  and  T.  I.  Storer.  Animal  Life  in  the  Yosemitc.  San  Francisco. 
Pp.  1-741. 

1925  Storer,  T.  I.  Univ.  Calif.  Pub.  ZooL,  27,  pp.  1-342. 

1928    Klauber,  L.  M.  Zool.  Soc.  San  Diego,  Bull.  no.  4,  pp.  1-8.  2d  ed.,  1930, 
no.  5. 

1928  Slevm,  J.  R.  (see  British  Columbia). 

1929  Burt,  C.  E.,  and  M.  D.  Burt  (see  Arizona). 

1929  Klauber,  L.  M.  Copeia,  no.  170,  pp.  15,  16. 

1930  Bogert,  C.  M.  Bull.  S.  Calif.  Acad.  Sci.,  29%  1-14. 

1930    Grinnell,  J.,  J.  Dixon,  and  J.  M.  Linsdale.  Univ.  Calif.  Pub.  ZooL,  35, 139- 

144. 

1930     Klauber,  L.  M.  (see  Klauber,  1928,  above). 
1930    Myers,  G.  S.  Copeia,  43,  61-64. 
1930    Myers,  G.  S.  Proc.  Btol.  Soc.  Wash.,  pp.  73-77. 

1930  Pick  well,  G.,  E.  Smith,  and  K.  Hazcltmc.  Western  Nature  Study  (San 
Jose),  i1,  1-55. 

1931  Myers,  G.  S.  Copeia,  no.  2,  pp.  56,  57. 

1932  Brues,  C.  T.  (see  Nevada). 

1932    Klauber,  L.  M.  Copeia,  no.  3,  pp.  118-120. 

1934    Eckert,  J.  E.  Copeia,  no.  2,  pp.  92-93. 

i934a    Klauber,  L.  M.  ZooL  Soc.  San  Diego,  Bull.  no.  n,  pp.  1-28. 

1936    Fitch,  H.  S.  (see  Oregon). 


596  HANDBOOK  OF  FROGS  AND  TOADS 

1936  Grinnell,  J.,  and  J.  M.  Linsdale.  Carnegie  Inst.  Pub.,  no.  481,  pp.  35-39. 

1936  Miller,  L.,  and  A.  H.  Miller.  Copeia,  no.  3,  p.  176. 

1937  Shapovalov,  L.  Copeia,  no.  4,  p.  234. 

1938  Schmidt,  K.  P.  (see  Illinois). 

1939  Wood,  W.  F.  Copeia,  p.  no. 

1940  Smith,  R.  E.  Science,  922391,  379,  380. 

1940  Stanton,  K.  Copeia,  no.  2,  p.  136. 

1941  Twitty,  V.  C.  Copeia,  no.  i,  pp.  1-3. 

1941  Vestal,  E.  H.  Copeia,  no.  3,  pp.  183. 

1942  Myers,  G.  S.  Occ.  Papers  Mus.  ZooL,  Stanford  Univ.,  no.  460,  pp.  1-13 
(plus  8  pp.  of  plates). 

1942  von  Bloeker,  J.  C.,  Jr.  Bull.  So.  Calif.  Acad.  Sci.,  41,  pt.  i,  29-38. 

1943  Marr,  J.  C.  Copeia,  no.  i,  p.  56. 

1943    Myers,  G.  S.  Copeia,  no.  2,  pp.  125-126. 

1943  Wiggins,  I.  L.  Copeia,  no.  3,  p.  197. 

1944  Miller,  R.  R.  Copeia,  no.  2,  p.  123. 

1944  Myers,  G.  S.  Copeia,  no.  i,  p.  58. 

1945  Pequegnat,  W.  E.  Jour.  Ent.  &  ZooL,  37%  1-7. 
Colorado 

1875    Yarrow,  H.  C.  (see  Arizona). 

1910  Cockerell,  T.  D.  A.  Univ.  Colo.  Studies,  82,  130. 

1911  Gary,  M.  North  Amer.  Fauna,  no.  33,  pp.  24,  27,  40. 

1913-1915    Ellis,  M.  M.,  and  J.  Henderson.  Univ.  Colo.  Studies,  io2,  pt.  i,  39-61 ; 
ii4,  pt.  2,  253-259. 

1924  Gilmore,  R.  J.  Colo.  Coll.  Pub.,  gen.  ser.  no.  129,  science  ser.,  13%  1-12. 

1926  Burnett,  W.  L.  Occ.  Papers  Mus.  ZooL  and  Ent.  (Colo.  State  Coll.),  i1, 
1-4. 

1927  Cockerell,  T.  D.  A.  Zoology  of  Colorado.  Boulder.  Pp.  112-113. 
i929b    Burt,  C.  E.,  and  M.  D.  Burt  (see  Arizona). 

19333    Burt,  C.  E.  (see  General  Worlds). 

1935  Burt,  C.  E.  (see  General  Worlds). 
Connecticut 

1844    Linsley,  J.  H.  Amer.  Jour.  Sci.  and  Arts,  ist  sen,  46,  47-48. 

1898    Sherwood,  W.  L.  Proc.  Linn.  Soc.  N.Y.  1897-1898,  no.  10,  pp.  9-24. 

1904  Henshaw,  S.  Fauna  of  New  England.  Occ.  Papers  Boston  Soc.  Nat.  Hist., 
72,  10-17. 

1905  Ditmars,  R.  L.  Amer.  Mus.  Jour.,  5%  184-206. 
1927     Babcock,  H.  L.  Copeia,  no.  90,  p.  8. 

1926  Babcock,  H.  L.  Bull.  Boston  Soc.  Nat.  Hist.,  no.  38,  pp.  11-14. 

1932  Babbitt,  L.  H.  Same,  no.  63,  pp.  25-26. 

1933  Ball,  S.  C.  Anat.  Rec.  47  (4  Suppl.),  p.  101. 

1936  Ball,  S.  C.  Trans.  Conn.  Acad.  Arts  Sci.,  32,  351-359. 

1937  Babbitt,  L.  H.  State  Geol.  and  Nat.  Hist.  Survey  (Conn.),  Bull.  no.  57. 
Hartford.  Pp.  50,  20  pis. 

Delaware 

1906  Stone,  W.  Amer.  Naturalist,  40, 162-164. 

1925  Fowler,  H.  W.  Copeia,  no.  145,  pp.  57-59. 


BIBLIOGRAPHY 


597 


1940  Conant,  R.  Herpetologica,  i,  176-177. 

1945  Conant,  R.  Soc.  Nat.  Hist.  Dela.  (Wilmington),  Feb.,  1945,  pp.  1-8. 

District  of  Columbia 

1899  Miller,  G.  S.,  Jr.  Proc.  BloL  Soc.  Wash.,  13,  75-78. 

1902  Hay,  W.  P.  Same,  15,  127-131. 

1915  Gronberger,  S.  M.  Copeia,  no.  24,  pp.  54-55. 

1918  Dunn,  E.  R.  (see  Virginia). 

1918  McAtee,  W.  L.  Bull.  Biol.  Soc.  Wash.,  no.  i,  pp.  44-46. 

19373  Brady,  M.  K.  (see  Maryland). 

i937b    Brady,  M.  K.  Proc.  Biol.  Soc.  Wash.t  50,  pp.  137-139. 
Florida 

1857  Hallowell,  E.  Proc.  Acad.  Nat.  Set.  Phila.  (1856),  8,  141,  142,  143. 

1871  Cope,  E.  D.,  2nd  and  yd  Ann.  Reft.  Peabody  Acad.  Salem.  Pp.  182. 

1877  Cope,  E.  D.  Amer.  Naturalist,  n9,  565. 

i877a  Cope,  E.  D.  Proc.  Amer.  Phil.  Soc.f  17,  87-88. 

1888  Cope,  E.  D.  Proc.  U.S.  Nat.  Mus.  (  1887),  10,  436. 

1895  Cope,  E.  D.  Proc.  Acad.  Nat.  Sci.  Phila.  (1894,  1895),  46,  429,  437,  438. 

1895  Loennberg,  E.  Proc.  U.S.  Nat.  Mus.  (1894),  17,  338-339. 

1901  Stejneger,  L.  Same,  24,  212-213. 

1905  Stejneger,  L.  In  G.  B.  Shattuck,  The  Bahama  Islands.  New  York.  Pp. 


1906  Fowler,  H.  W.  Proc.  Acad.  Nat.  Sci.  Phila.,  58,  109. 

1910  Barbour,  T.  Proc.  Biol.  Soc.  Wash.,  23,  100. 

1910  Brimley,  C.  S.  (see  Alabama). 

1910  Barbour,  T.  Mem.  Mus.  Comp.  ZooL,  44%  209-359. 

1914  Deckert,  R.  F.  Copeia,  no.  5,  pp.  2-4;  no.  9,  pp.  1-3. 

1915  Deckert,  R.  F.  Copeia,  no.  18,  pp.  3-5. 
19153  Deckert,  R.  F.  Copeia,  no.  20,  pp.  21-22. 
1917  Fowler,  H.  W.  Copeia,  no.  43,  p.  39. 

1920  Barbour,  T.  Copeia,  no.  84,  pp.  55-56. 

1921  Deckert,  R.  F.  Copeia,  no.  92,  pp.  20-23. 

1922  Deckert,  R.  F.  Copeia,  no.  112,  p.  88. 

1923  Hallinan,  T.  Copeia,  no.  115,  pp.  19,  20. 
1923  Van  Hyning,  T.  Copeia,  no.  118,  p.  68. 
1926  Dunn,  E.  R.  Copeia,  no.  157,  pp.  154-156. 
1931  Barbour,  T.  Copeia,  no.  3,  p.  140. 

1933  Van  Hyning,  O.  C.  Copeia,  no.  i,  pp.  5-7. 

1934  Carr,  A.  F.,  Jr.  Fla.  Naturalist,  72,  19-23. 

1935  Barbour,  T.  Zoologica,  19%  78,  79,  89-91,  96. 

1935  Brady,  M.  K.,  and  F.  Harper.  Proc.  Biol.  Soc.  Wash.,  48,  107-1  10. 

1935  Harper,  F.  Proc.  Biol.  Soc.  Wash.,  48,  79-82. 

1935  Harper,  F.  (see  General  Wor^s). 

1938  Allen,  E.  R.  Copeia,  no.  i,  p.  50. 

1938  Goin,  C.  J.  Copeia,  no.  i,  p.  48. 

1938  Springer,  S.  Copeia,  no.  i,  p.  49. 

1939  Allen,  E.  R.  Copeia,  no.  i,  p.  53. 

Campbell,  G.  R.,  and  W.  H.  Stickel.  Copeia,  no.  2,  p.  105. 


598  HANDBOOK  OF  FROGS  AND  TOADS 

1939  Harper,  F.  (see  Georgia). 

1939  Skermer,  G.  A.  Copeia,  no.  2,  pp.  107-108. 

1940  Allen,  [E.]  R.,  and  M.  P.  Merryday.  Nat.  Hist.,  46 l,  234. 

1940  Blair,  A.  P.,  C.  C.  Hargreaves,  and  K.  K.  Chen.  Proc.  Soc.  Exp.  Biol.  Ued., 

45l,  209-214. 

i94oa  Carr,  A.  F.,  Jr.  Copeia,  no.  i,  p.  55. 

i94ob    Carr,  A.  F.,  Jr.  Univ.  Fla.  Pub.  Biol.  Sci.,  ser.  3,  no.  i,  pp.  1-118. 

1940  Goin,  C.  J.,  and  M.  G.  Netting  (see  I^ouisiana). 

1940  Stejneger,  L.  Copeia,  no,  3,  pp.  149-151. 

1941  Moore,  J.  A.  (see  General  Worths). 

1943  Goin,  C.  J.  Proc.  Fla.  Acad.  Sci.,  6 l* 4, 148-149. 

1944  Goin,  C.  J.  Copeia,  no.  3,  p.  192. 

1945  Allen,  E.  [R.  |,  and  R.  Slatten.  Hcrpetologica,  3,  pt.  i,  25. 
1945  Kilby,  J.  D.  Quart.  Jour.  Fla.  Acad.  Sci.,  8,  no.  i,  71-104. 
1945  Myers,  G.  S.  Copeia,  no.  i,  p.  44. 

1945  Netting,  M.  G.,  and  C.  J.  Goin.  Proc.  Fla.  Acad.  Sci.  (1944),  72~3>  181-184. 

1945  Shreve,  B.  Copeia,  no.  2,  p.  117. 

1947  Goin,  C.  }.  Univ.  Fla.  Studies,  Biol.  Sci.,  ser.  4,  no.  2,  1-66. 
Georgia 

1849  Holbrook,  }.  E.  In  George  White,  Statistics  of  Georgia.  Savannah.  App., 

p.  15. 

1857  Le  Conte,  J.  Proc.  Acad.  Nat.  Sci.  Phila.  (1856),  8,  146. 

1901  Fountain,  P.  The  Great  Deserts  and  Forests  of  North  America.  London, 

New  York,  and  Bombay.  Pp.  62-64. 

1908  Allard,  H.  A.  Science  (n.s.)9  2872n,  655-656. 

1910  Brimley,  C.  S.  (see  Alabama). 

1923  Wright,  A.  H.  Copeia,  no.  115,  p.  34. 

1926  Wright,  A.  H.  Ecology,  j\  81-83. 

1930  Harper,  F.  Copeia,  pp.  152-154. 
i93ia    Harper,  F.  Copeia,  pp.  159-161. 

1931  Harper,  F.  Sci.  Monthly,  32,  176-181. 

1932  Wright,  A.  H.  (see  General  Wor^s). 

1933  Brandt,  B.  B.,  and  C.  F.  Walker  (see  North  Carolina). 

1934  Carter,  H.  A.  Copeia,  no.  3,  p.  138. 

1935  Harper,  F.  (see  Florida). 

1937  Harper,  F.  Amer.  Midi.  Naturalist,  i82,  260-272. 

19393    Harper,  F.  Same,  22,  134-149. 

i939b    Harper,  F.  Notulae  Naturae  (Phila.),  27,  pp.  1-4. 

1942  Stewart,  L.  Nature  Magazine,  yf,  128-130. 

1948  Neill,  W.  T.  Herpetologica,  4%  108-109. 
1948  Neill,  W.  T.  Same,  4%  158. 

1948  Neill,  W.  T.  Same,  4-%  175-179. 
Idaho 

1869  Cooper,  J.  G.  (see  Montana). 

1872  Cope,  E.  D.  (see  Montana). 

1884  Cope,  E.  D.  (see  California). 

1884  Cope,  E.  D.  Proc.  Acad.  Nat.  Sci.  Phtla.  (1883),  35,  17. 


BIBLIOGRAPHY  599 

1891  Stejneger,  L.  North  American  Fauna,  no.  5,  pp.  112-113. 
1895    Van  Denburgh,  J.  Bull.  U.S.  Fish  Commission,  14, 207. 

1912    Van  Denburgh,  J.  Proc.  Calif.  Acad.  Sci.f  4th  ser.,  3,  158,  159. 
I92ib    Van  Denburgh,  J.,  and  J.  R.  Slevin.  Same,  n3,  39,  42. 

1928  Slevin,  J.  R.  (see  British  Columbia). 
1933    Linsdale,  J.  M.  Copeia,  no.  4,  p.  223. 

1940  Tanner,  W.  W.  (see  Utah). 

19413    Slater,  J.  R.  Occ.  Papers  Dept.  Biol.,  Coll.  Puget  Sound  (Tacoma),  no. 
14,  pp.  78-108. 

1941  Tanner,  W.  W.  Great  Basin  Naturalist,  2J,  87-90. 

1942  Slater,  J.  R.  (see  Washington). 
1944    Knowlton,  G.  F.  (see  Utah). 

1946    Evenden,  F.  G.  Copeia,  no.  4,  p.  257. 
Illinois 

1883     Davis,  N.  S.,  Jr.,  and  F.  L.  Rice.  Bull.  Chicago  Acad.  Sciences,  i\  25,  27, 
28. 

1889  Roberts,  H.  L.  Amer.  Naturalist,  23-%  74. 

i89oa    Garman,  H.  Bull.  III.  State  Lab.  Nat.  Hist.,  3, 133,  134. 

1890  Garman,  H.  Same,  3,  188-190. 

1892  Garman,  H.  Same,  3,  art.  XIII,  316-352. 

1892  Hay,  O.  P.  (see  Indiana). 

1893  Hurter,  J.  Trans.  Acad.  Sci.  St.  Louis,  6M,  251-254. 
1902    Garman,  H.  Bull.  111.  State  Lab.  Nat.  Hist.,  3,  335-339- 

1914  Gaige,  H.  T.  Copeia,  no.  IT,  p.  4. 

1915  Thompson,  C.  Occ.  Papers  Mus.  Zool.  Univ.  Mich.,  9,  1-7. 

1917  Hankmson,  T.  L.  Trans.  III.  Acad.  Set.,  Tenth  Ann.  Meeting,  10,  pp. 
324-325. 

1918  Hubbs,  C.  L.  Copeia,  no.  55,  pp.  40-43. 

1919  Pope,  P.  H.  Copeia,  no.  74,  pp.  83,  84. 

1923  Weed,  A.  C.  Copeia,  no.  116,  pp.  45-46,  48,  49. 

1924  Ridgway,  R.  Copeia,  no.  128,  p.  39. 

1925  Blanchard,  F.  N.  Papers  Mich.  Acad.  Sci.  Arts  Letters,  pp.  534-535- 

1926  Cahn,  A.  R.  Copeia,  no.  151,  pp.  107-109. 
19293    Burl,  C.  E.,  and  M.  D.  Burt  (see  Arkansas). 
I929b    Burt,  C.  E.,  and  M.  D.  Burt  (see  Arizona). 

1929  Schmidt,  K.  P.  Field  Museum  Nat.  Hist.  Zool.  Leaflet,  no.  n,  pp.  1-15. 
1933  Davis,  D.  D.  Copeia,  no.  4,  pp.  223-224. 

1935    Burt,  C.  E.  (see  General  Wor\s). 

1935    Schmidt,  K.  P.,  and  W.  L.  Necker.  Bull.  Chicago  Acad.  Sci.,  5*,  57-61,  64- 

67,  76-77. 
1938    Necker,  W.  L.  Chicago  Acad.  Sci.,  Leaflet  no.  i,  p.  2. 

1938  Schmidt,  K.  P.  Field  Mus.  Nat.  Hist.,  Zool.  ser.,  2o20,  377-382. 

1939  Necker,  W.  L.  Bull.  Chicago  Acad.  Sci.,  61,  i,  2,  4-6. 

1941     Cagle,  F.  R.  Cont.  So.  III.  Norm.  Univ.,  Mus.  Nat.  and  Soc.  Sci.  Car- 
bondale.  IV.  32  pp. 

1941  Owens,  D.  W.  Copeia,  No.  3,  pp.  183,  184. 

1942  Cagle,  F.  R.  Amer.  Midi.  'Naturalist,  28*, 164-200. 


6oo  HANDBOOK  OF  FROGS  AND  TOADS 

1942    Gloyd,  H.  K.  Trans.  Ill  Acad.  Set.,  34%  220. 
1942    Peters,  J.  A.  Copeia f  no.  3,  p.  182. 

1944  Pope,  C.  L.  Amphibians  and  Reptiles  of  the  Chicago  Area.  Nat.  Hist. 
Mus.,  Chicago.  Pp.  1-275. 

1945  Elder,  W.  H.  Copeia,  no.  2,  p.  122. 

1946  Conant,  R.  Copeia,  no.  2,  p.  109. 
Indiana 

1886  Hughes,  E.  Brootyille  Soc.  Nat.  Hist.,  no.  2,  p.  43. 

1887  Butler,  A.  W.  Jour.  Cine.  Soc.  Nat.  Hist.,  io;i,  147-148. 
1887    Hay,  O.  P.  Same,  io2,  59,  62-63. 

1887    James,  J.  E.  (see  Ohio}. 

1889    Hay,  O.  P.  Amer.  Naturalist,  23<J,  770-774. 

1891  Blatchley,  W.  S.  Jour.  Cincinnati  Soc.  Nat.,  14,  27-28. 
1891-1892    Butler,  A.  W.  Same,  13,  175. 

1892  Hay,  O.  P.  Ind.  Deft.  Geol.  and  Nat.  Res.,  i7th  Ann.  Rept.  (1891),  pp. 
456-481,  585,  586. 

18953    Kirsch,  P.  H.  Bull.  U.S.  Fish  Commission  (1894),  14,  art.  6,  41. 

iSgsb    Kirsch,  P.  H.  Same,  art.  20,  p.  333. 

1896    Atkinson,  C.  Proc.  Ind.  Acad.  Set.  (1895),  pp.  258-261. 

1900  Blatchley,  W.  L.  Ind.  Dept.  Geol.  and  Nat.  Res.,  24th  Ann.  Rept.  (1899), 
pp.  542,  543,  551. 

1901  Ramsey,  E.  E.  Proc.  Ind.  Acad.  Sci.  ( 1900),  p.  222. 
1907    Banta,  A.  M.  Carnegie  Inst.  Pub.  Washington. 
1907    McAtee,  W.  L.  Proc.  BioL  Soc.  Wash.,  20,  15,  16. 

1909    Hahn,  W.  L.  Proc.  U.S.  Nat.  Mus.,  35  (Dec.  7,  1908),  545-548,  557-562. 
1918    Hubbs,  C.  L.  (see  Illinois). 

1920  Evermann,  B.  W.,  and  H.  W.  Clark.  La^e  Maxintyicfye,  Ind.  Dept. 
Conservation  Pub.,  i,  631-644. 

1921  Ortenburger,  A.  I.  Copeia,  no.  99,  pp.  73-75. 

1926    Blanchard,  F.  N.  Papers  Mich.  Acad.  Sci.  Arts  Letters  (1925),  pp.  371- 

373- 

1926  Myers,  G.  S.  Proc.  Ind.  Acad.  Sci.  (1925),  35,  281-286. 

Myers,  G.  S.  Proc.  Ind.  Acad.  Sci.f  36,  338-339. 
Myers,  G.  S.  Copeia,  no.  163,  pp.  50-53. 

1927  Wright,  H.  P.,  and  G.  S.  Myers,  Copeia,  no.  159,  pp.  173-175. 

1928  Springer,  S.  Proc.  Ind.  Acad.  Sci.  (1927),  37,  492. 
1931     Grave,  B.  H.,  Proc.  Ind.  Acad.  Sci.  (1930),  40,  339. 
1931    Piatt,  J.  Same,  pp.  362,  363,  365. 

1935  Burt,  C.  E.  (see  General  Worlds). 

1935  Schmidt,  K.  P.,  and  W.  L.  Necker  (see  Illinois). 

1936  Grant,  C,  Proc.  Ind.  Acad.  Sci.,  45,  323-333. 
1938  Necker,  W.  L.  (see  Illinois). 

1938    Schmidt,  K.  P.  (see  Illinois). 

1938  Swanson,  P.  L.  Amer.  Midi.  Naturalist,  2O3,  713. 

1939  Miller,  D.  C.  Proc.  Ind.  Acad.  Sci.  (1940),  49,  209-214. 

1939  Necker,  W.  L.  (see  Illinois). 

1940  Swanson,  Paul  L.  Amer.  Midi.  Naturalist,  22%  684-695. 


BIBLIOGRAPHY  601 

19413    Blair,  A.  P.  Proc.  Nat.  Acad.  Sci.,  27*,  14-17. 

1942  Vogel,  H.  H.,  Jr.  Proc.  Ind.  Acad.  Sci,  51,  p.  266. 
Iowa 

1878    Aldrich,  C.  Amer.  Naturalist,  I27,  473-474. 

1892    Osborn,  H.  A  Partial  Catalogue  of  the  Animals  of  Iowa.  Ames.  P.  10. 
i9iob    Ruthven,  A.  G.  Proc.  Iowa  Acad.  Sci.  (1910),  17,  198-205. 
19126    Ruthven,  A.  G.  Same  (1912),  19,  207. 

1919    Ruthven,  A.  G.  Occ.  Papers  Mus.  Zool.  Univ.  Mich.,  no.  66,  pp.  1-2. 
1923     Blanchard,  F.  N.  Univ.  Iowa  Studies  in  Nat.  Hist.,  new  ser.  no.  67,  io2, 
19-22. 

Burt,  C  E.,  and  M.  D.  Burt  (see  Arkansas). 

Burt,  C.  E.,  and  M.  D.  Burt  (see  Arizona). 

1935  Burt,  C.  E.  (see  General  Worlds). 

1941     Bailey,  R.  M.,  and  M.  K.  Bailey.  Iowa  State  Coll.  Jour.  Sci.t  15,  169-177. 

1943  Bailey,  R.  M.  Iowa  Acad.  Sci.t  50,  347-352. 

1944  Bailey,  R.  M.  Iowa  Conservationist,  3%  17-20;  3*,  25,  27-30. 

1945  Abbott,  R.  L.  Nat.  Hist.,  54%  114-117. 
1948    Loomis,  R.  B.  Herpetologica,  4*,  121-122. 

Kansas 

18573  Hallowell,  E.  Proc.  Acad.  Nat.  Sci.  Phila.  (1856),  8, 250-252. 

i857b  Hallowell,  E.  (see  Texas). 

1881  Cragm,  F.  W.  Trans.  Kans.  Acad.  Sci.,  7,  121-123. 

1885  Cragin,  F.  W.  Same,  9,  136-140. 

1894  Cragin,  F.  W.  Colo.  Coll.  Studies,  5,  39. 

1907  Hartmann,  F.  A.  Trans.  Kans.  Acad.  Sci.  (1906),  20,  pt.  2,  227-229. 

1925  Dice,  L.  R.  Ecology,  4  (Jan.,  1923),  40-53. 

1927  Burt,  C.  E.  Occ.  Papers  Mus.  Zool.  Univ.  Mich.,  no.  189,  pp.  1-3. 

1927  Linsdale,  J.  M.  Copeia,  no.  164,  pp.  75-77. 

1928  Gloyd,  H.  K.  Trans.  Kans.  Acad.  Sci.,  31,  pp.  116-119. 
19293  Burt,  C.  E.,  and  M.  D.  Burt  (see  Arkansas). 

T92c;b    Burt,  C.  E.,  and  M.  D.  Burt  (see  Arizona). 

1929  Gloyd,  H.  K.  Science,  691776,  44. 

i929b     Taylor,  E.  H.  Kans.  Univ.  Sci.  Bull.,  19°,  65. 

1931  Burt,  C.  E.  Proc.  Biol.  Soc.  Wash.,  44,  1 1-14. 

1932  Gloyd,  H.  K.  Papers  Mich.  Acad.  Sci.  (1931),  15,  389-400. 

1932  Smith,  H.  M.  Trans.  Kans.  Acad.  Sci.,  35,  93-96. 

1933  Smith,  H.  M.  Copeia,  no.  4,  p.  217. 

1934  Brennan,  L.  A.  Trans.  Kans.  Acad.  Sci.t  37, 189-191. 

1934    Smith,  H.  M.  Amer.  Midi.  Naturalist,  15*,  377-390,  427-528. 

1936  Hibbard,  C.  W.,  and  A.  B.  Leonard.  Copeia,  no.  2,  p.  114. 

1936  Youngstrom,  K.  A.,  and  H.  M.  Smith.  Amer.  Midi.  Naturalist,  17*, 
629-633. 

1937  Brenkelman,  J.  P.,  and  A.  Downs.  Trans.  Kans.  Acad.  Sci.,  39,  267-268. 

1937  Grant,  C.  Amer.  Midi.  Naturalist,  18,  370-372. 

1938  Brennan,  L.  A.  Trans.  Kans.  Acad.  Sci.,  40,  341-347. 

1938  Tihen,  J.  A.  Same,  pp.  401-409. 

1939  Tihen,  J.  A.,  and  J.  M.  Sprague.  Same,  42,  499-512. 


602  HANDBOOK  OF  FROGS  AND  TOADS 

1944  Marr,  J.  C.  (see  General  Worlds). 

1945  Mittleman,  M.  B.  Herpetologica,  3,  pt.  i,  20—21. 

1946  Breukelman,  J.,  and  H.  M.  Smith.  Univ.  Kans.  Pub.  Mus.  Nat.  Hist.,  i5, 
103-105. 

1947  Smith,  H.  M.  Herpetologica,  4,  13-14. 
Kentucky 

1894    Garman,  H.  Bull.  Essex  Inst.,  26,  36-37. 

1901    Garman,  H.  Ky.  Ag.  Expt.  Stat.  Bull.,  no.  91,  pp.  62-64. 

1926    Bishop,  S.  C.  Copeia,  no.  152,  pp.  118-120. 

1926    Blancharcl,  F.  N.  (see  Indiana). 

19293     Burt,  C.  E.,  and  M.  D.  Burt  (see  Arkansas). 

i933b    Burt,  C.  E.  Amer.  Midi.  Naturalist,  14°,  669-679. 

1934  Bailey,  V.  Same  (1933),  M5,  594-596- 

1935  Harper,  F.  (sec  General  Worlds). 

1936  Giovanolh,  L.  Copeia,  no.  i,  p.  69. 

1937  Hibbard,  C.  W.  Trans.  Kans.  Acad.  Sci.,  39,  277-281. 
1939    Parker,  M.  V.  (sec  Tennessee}. 

1939  Welter,  W.  A.,  and  K.  Carr.  Copeia,  no.  3,  pp.  128-130. 

1940  Dury,  R.,  and  W.  Gessing,  Jr.  Herpetologica,  2,  pt.  2,  31,  32. 

1941  Barbour,  R.  W.,  and  E.  P.  Walters.  Copeia,  no.  2,  p.  116. 

1941  Barbour,  R.  W.  Copeia,  no.  4,  p.  262. 

1942  Barbour,  R.  W.  Copeia,  no.  2,  p.  128. 
1946    Barbour,  R.  W.  Copeia,  no.  r,  p.  44. 

Louisiana 

1900    Beyer,  G.  E.  Proc.  La.  Soc.  Naturalists  (1897-1899),  Appendix,  i,  35-37. 
1918    Viosca,  Percy,  Jr.  Btenn.  Rept.  La.  Dept.  Conservation  (Apr.  i,  1916), 

pp.  160-162. 

1920     Schmidt,  K.  P.  Copeia,  no.  86,  pp.  84—85. 
19233    Viosca,  P.,  Jr.  Copeia,  no.  115,  pp.  35-44- 
i923b    Viosca,  P.,  Jr.  Copeia,  no.  122,  pp.  96-99. 
1926    Strecker,  J.  K.,  Jr.,  and  L.  S.  Frierson,  Jr.  Cont.  Baylor  Univ.  Mus.,  no.  5, 

pp.  4-5. 

1926    Viosca,  P.,  Jr.  Ecology,  7,  307-314. 
1928    Viosca,  P.,  Jr.  Proc.  Biol.  Soc.  Wash.,  41,  89-92. 
i92Qa    Burt,  C.  E.,  and  M.  D.  Burt  (see  Arkansas}. 
1935     Burt,  C.  E.  (see  General  Worlds). 
1935    Harper,  F.  (see  General  Worlds). 

1938  George,  I.  D.  Proc.  La.  Acad.  Sci.,  4,  255-259. 

1939  Schmidt,  K.  P.  (see  Illinois). 

1939  La.  Dept.  Conservation,  Div.  Fisheries,  Bull.  26. 

1940  Blair,  A.  P.,  C.  C.  Hargreavcs,  and  K.  K.  Chen  (see  Florida). 
1940    George,  I.  D.  Copeia,  no.  2,  p.  134. 

1940  Coin,  C.  J.,  and  M.  Graham  Netting.  Ann.  Carnegie  Mus.,  28,  137-168. 

1942  Palmer,  M.  E.  Proc.  La.  Acad.  Sci.,  6,  73-74. 

1944  Marr,  J.  C.  (see  General  WorJ(s). 

1944  Viosca,  P.,  Jr.  Proc.  La.  Acad.  Sci.t  8,  47-62. 

1945  Burt,  C.  E.  Trans.  Kans.  Acad.  Sci.  (1945-1946),  483,  423-425. 


BIBLIOGRAPHY  603 

1947    Orton,  G.  L.  Ann  Carnegie  Mus.  30,  363-383. 
Maine 

1832  Williamson,  W.  D.  History  of  the  State  of  Maine.  Hallowell.  Vol.  I,  sect. 
V,  p.  169. 

1862  Fogg,  B.  F.  Proc.  Portland  Soc.  Nat.  Hist.,  i,  pt.  I,  86. 

1863  Verrill,  A.  E.  Proc.  Boston  Soc.  Nat.  Hist.  (1862,  1863),  9,  197-198.  Bos- 
ton, 1865.  (Read  Feb.  4,  1863.) 

1873  Adams,  A.  L.  (see  New  Brunswic^). 

1873  Hill,  T.  Amer.  Naturalist,  7",  660-662. 

1904  Henshaw,  S.  (see  Connecticut). 

1915  Pope,  P.  H.  Copeia,  no.  16,  pp.  1-2. 

1918  Pope,  P.  H.  Copeia,  no.  64,  pp.  96-97. 

1926  Babcock,  H.  L.  (see  Connecticut). 

1938  Hoopes,  I.  New  Eng.  Naturalist,  no.  T,  pp.  4-6. 

1939  Manville,  R.  H.  Copeia,  p.  174. 

1942  Fowler,  J.  A.  Copeia,  no.  3,  pp.  185-186. 
Maryland 

1902  Hay,  W.  P.  (see  District  of  Columbia). 

1914  Keim,  T.  D.  Copeia,  no.  2,  p.  2. 

1915  Fowler,  H.  W.  Copeia,  no.  22,  pp.  38-39. 
1918  McAtee,  W.  L.  (see  District  of  Columbia). 
1925  Fowler,  H.  W.  Copeia,  no.  145,  pp.  61-63. 
1932  Walker,  C.  F.  (see  Ohio). 

1936  Noble,  G.  K.,  and  W.  G.  Hassler.  Copeia,  no.  i,  pp.  63-64. 
i937a    Brady,  M.  K.,  Proc.  Biol.  Soc.  Wash.,  50, 190-191. 

i937b    Brady,  M.  K.  Same,  pp.  137-138. 

1937  Kelly,  H.  A.  Snakes  of  Maryland,  Baltimore.  Pp.  1-103. 

1939  Robertson,  H.  C.  Bull.  Nat.  Hist.  Soc.  Md.,  910,  89-93. 

1940  McCauley,  R.  H.,  Jr.,  and  C.  S.  East.  Copeia,  no.  2,  pp.  120-123. 

1940  Mansueti,  R.  Bull.  Nat.  Hist.  Soc.  Md.,  io10,  88-96. 

1941  Mansueti,  R.  Proc.  Nat.  Hist.  Soc.  Md.,  7, 1-53. 

1942  Mansueti,  R.  Bull.  Nat.  Hist.  Soc.  Md.,  iaa,  33-43. 

1946  Littleford,  R.  A.  Copeia,  no.  2,  p.  104. 

1947  Fowler,  J.  A.,  Maryland,  17,  6,  7. 
Massachusetts 

1833  Smith,  D.  S.  C.  H.  In  Kept.  Geol.  Mm.  Bot.  Zool,  by  E.  Hitchcock,  pt. 

IV,  p.  552. 

1839  Storer,  D.  H.  Repts.  Mass.  Commissioners  1838,  Zool.  Survey.  Boston. 
Pp.  235-245. 

1840  Storer,  D.  H.  Boston  Jour.  Nat.  Hist.,  31  2,  40-53. 

1852    Nichols,  A.  Jour.  Essex  Co.  Nat.  Hist.  Soc.,  i8,  113-117.  (Read  June  17, 

1853.) 
1865    Putnam,  F.  W.  Proc.  Boston  Soc.  Nat.  Hist.,  9,  229-230. 

1867  Putnam,  F.  W.  Amer.  Naturalist,  i,  107-109. 

1868  Allen,  J.  A.  Proc.  Boston  Soc.  Nat.  Hist.,  12,  185-198,  249. 
1870    Allen,  J.  A.  Same,  13  (1869-1871),  261,  263.  March  16,  1870. 
1873    Fowler,  S.  P.  Amer.  Naturalist,  7*,  239. 


604  HANDBOOK  OF  FROGS  AND  TOADS 

1880    Hinckley,  M.  H.  Proc.  Boston  Soc.  Nat.  Hist.,  21  (1880-1882),  104-107. 
Nov.  17,  1880. 

1882  Hinckley,  M.  H.  Amer.  Naturalist,  16,  636-639. 

1883  Hinckley,  M.  H.  Proc.  Boston  Soc.  Hist.,  21, 307-315. 

1884  Hinckley,  M.  H.  Same  ( 1882-1883),  22,  85-95. 
1888    Hargett,  C.  U.  Amer.  Naturalist,  22,  536,  537. 

1897    Kirkland,  A.  H.  Hatch  Expt.  Stat.,  Mass.  Ag.  Coll.  Bull.,  no.  46,  pp.  1-29. 
1904    Henshaw,  S.  (see  Connecticut). 
1926    Babcock,  H.  L.  (see  Connecticut). 
1930    Dunn,  E.  R.  Bull.  Boston  Soc.  Nat.  Hist.,  no.  57,  pp.  5-6. 
1936    Driver,  E.  G.  Copeia,  no.  i,  pp.  67,  68,  69. 
1941     Moore,  J.  A.  (see  General  Worlds). 
1944    Sweetman,  H.  L.  Amer.  Midi.  Naturalist,  yr,  499-501. 
1947    Blair,  A.  P.  (see  New  Jersey). 
1947    Hoopes,  I.  Copeia,  no.  2,  pp.  138-139. 
Michigan 

1861     Miles,  M.  Geol.  Survey  Mich.,  First  Bienn.  Kept.,  p.  234. 

1865    Cope,  E.  D.  Proc.  Acad.  Nat.  Sci.  Phila.,  17,  84-85. 

1879    Smith,  W.  H.  Science  News,  Supplement,  i23,  i,  v,  viii. 

1895    Gibbs,  M.  Amer.  Field,  pp.  179,  203,  228,  251,  273,  274,  299,  325. 

i895b    Kirsch,  P.  H.  (see  Indiana). 

1904    Clark,  H.  L.  Mich.  Acad.  Sci.,  Fourth  Ann.  Kept.  (1902),  p.  192. 

1906    Ruthven,  A.  G.  Geol.  Survey  Mich.,  Ann.  Rept.,  1905,  pp.  109-110. 

1908  Hankinson,  T.  L.  State  Board  Geol.  Survey  Mich.  1907-1908,  Rept., 
PP-  235-236. 

1909  Ruthven,  A.  G.  Geol.  Survey  Mich.,  Rept.,  pp.  329-333. 
I909a    Ruthven,  A.  G.  Mich.  Acad.  Sci.,  Eleventh  Rept.,  p.  116. 

1910  Ruthven,  A.  G.  Same,  Rept.,  12,  59. 

19113    Ruthven,  A.  G.  Mich.  Geol.  Biol.  Survey  Pub.,  4,  Biol.  ser.  2,  pp.  257- 

263. 

191  ib    Ruthven,  A.  G.  Mich.  Acad.  Set.,  Thirteenth  Rept.,  p.  115. 

1911  Thompson,  C.  Same,  pp.  105—106. 

1912  Ruthven,  A.  G.,  C.  Thompson,  and  H.  Thompson.  Mich.  Geol.  Biol. 
Survey  Pub.,  10,  Biol.  sen,  pp.  1-166. 

1912    Thompson,  C.,  and  H.  Thompson.  Mich.  Acad.  Sci.,  Fourteenth  Rept., 

pp.  156-157. 
1912    Thompson,  H,  Same,  p.  189. 

1912  Thompson,  C.  Same,  p.  190. 

1913  Thompson,  C.,  and  H.  Thompson.  Same,  Fifteenth  Rept.,  pp.  215-216. 

Gaige,  H.  T.  Copeia,  no.  14,  p.  2. 

Gaige,  H.  T.  Occ.  Papers  Univ.  Mus.  Univ.  Mich.,  no.  17,  pp.  1-4. 

1915  Thompson,  C.  Occ.  Papers  Mus.  Zool.  Univ.  Mich.,  no.  18,  pp.  1-4. 
19153    Thompson,  C.  Same,  no.  9,  pp.  1-7. 

1916  Thompson,  C.  Mich.  Geol.  Biol.  Survey  Pub.,  20,  Biol.  ser.  4,  pp.  62,  63. 
iQi6    Evans,  A.  T.  Proc.  U.S.  Nat.  Mus.,  49,  351-354. 

1917  Ellis,  M.  M.  Mich.  Acad.  Sci.,  Nineteenth  Rept.,  pp.  45-48. 

1917    Ruthven,  A.  G.  Occ.  Papers  Mus.  Zool.  Univ.  Mich.,  no.  47,  pp.  1-5. 


BIBLIOGRAPHY  605 

1918  Hubbs,  C.  L.  (see  Illinois). 

1920  Potter,  D.  Copeia,  no.  82,  pp.  89-90. 

1924  Hatt,  R.  T.  Papers  Mich.  Acad.  Set.,  pp.  374,  385. 
1928  Blanchard,  F.  N.  Copeia,  no.  167,  pp.  45-47. 
19290.    Burt,  C.  E.,  and  M.  D.  Burt  (see  Arkansas). 
1933  Blanchard,  F.  N.  Copeia,  no.  4,  p.  216. 

1933  Force,  E.  R.  Copeia,  no.  3,  pp.  128-13 1. 

1935  Schmidt,  K.  P.,  and  W.  L.  Necker  (see  Illinois). 

1937  Allen,  D.  Copeia,  no.  19,  no.  3,  pp.  190,  191. 

1938  Allen,  D.  L.  Ecol.  Monog.,  83,  430,  431. 

1938  Necker,  W.  L.  (see  Illinois). 

1939  Necker,  W.  L.  (see  Illinois). 

1940  George,  I.  D.  (see  Louisiana). 

1941  Moore,  J.  A.  (see  General  Worf(s). 

1942  Edgren,  R.  A.,  Jr.  Copeia,  no.  3,  p.  180. 

1944  Greaser,  C.  W.  Papers  Mich.  Acad.  Sci.  Arts  Letters  (1943),  29,  229—249 

(esp.  234-236). 
Minnesota 

1922  Weed,  A.  C.  Proc.  Biol.  Soc.  Wash.,  35,  107-110. 

1923  Weed,  A.  C.  Copeia,  p.  28. 
1930  Weed,  A.  C.  Turtox  News,  8,  43. 

1935  Swanson,  C}.  Copeia,  no.  3,  pp.  152-154. 

1938  Breckenndge,  W.  }.  Copeia,  no.  i,  p.  47. 

1941  Moore,  J.  A.  (see  General  Worlds) . 

1942  Moore,  J.  A.  Genetics,  27,  408-416. 

1944  Breckenridge,  W.  J.  Minn.  Mus.  Nat.  Hist.  Univ.  Minn.,  Univ.  Minn. 

Press.  Pp.  1-202. 
Mississippi 

1901  Stejneger,  L.  (see  Florida). 

1910  Brimley,  C.  S.  (see  Alabama). 

1920  Potter,  D.  Copeia,  no.  82,  p.  82. 

1927  Corrington,  }.  D.  Copeia,  no.  165,  pp.  98-100,  102. 
i932a    Allen,  M.  J.  Amer.  Mus.  Novitates,  no.  542,  pp.  1-2,  6-ir. 
i932b    Allen,  M.  J.  Copeia,  no.  2,  p.  104. 

1935  Harper,  F.  (see  General  Worlds). 

1938  Springer,  S.  (see  Florida) . 

1940  Coin,  C.  J.,  and  M.  G.  Netting  (sec  Louisiana). 

1942  Netting,  M.  G.,  and  C.  J.  Coin  (see  Alabama). 
Missouri 

1893  Hurter,  J.  (see  Illinois). 

1894  Test,  F.  C.  Bull.  U.S.  Fish  Commission,  12  (1892,  1894),  122. 
1897  Hurter,  J.  Trans.  Acad.  Sci.  St.  Louis,  7,  503. 

1903  Hurter,  J.  Same,  13, 80. 

1911  Hurter,  J.  Same,  20,  96-127. 

1921  Danforth,  C.  H.  Same,  24**. 

1925  Blanchard,  F.  N.  (see  Illinois). 

1929*    Burt,  C.  E.,  and  M.  D.  Burt  (see  Arkansas). 


606  HANDBOOK  OF  FROGS  AND  TOADS 

1933  Burt,  C.  E.  Amer.  Midi.  Naturalist,  I42,  170-173. 

1934  Boycr,  D.  A.,  and  A.  A.  Heinze.  Trans.  Acad.  Set.  St.  Louis,  28s"4,  185- 
200. 

1935  Burt,  C.  E.  (see  General  Worlds). 

1938    Henning,  W.  L.  Copeia,  no.  2,  pp.  91-92. 

1942    Anderson,  P.  Bull.  Chicago  Acad.  Sci.,  611,  203,  205-207. 

1945  Anderson,  P.  Same,  7%  272-273. 

1946  Peter,  J.  A.  Copeia,  no.  i,  p.  44. 
Montana 

1869    Cooper,  J.  G.  Amer.  Naturalist,  3,  125. 

1872    Cope,  E.  D.  In  U.S.  Geol.  Survey  of  Mont,  and  Adjacent  Territories, 
Prelim.  Kept.  (1871),  by  F.  V.  Hayden.  Pp.  468-469. 

1874  Allen,  J.  A.  Proc.  Boston  Soc.  Nat.  Hist.,  17,  70. 

1875  Hayden,  F.  V.  (see  Nebraska). 

1878  Coues,  E.,  and  H.  C.  Yarrow.  Bull.  U.S.  Geol.  and  Geog.  Survey,  4,  art. 
XI,  288-290. 

1879  Cope,  E.  D.  Amer.  Naturalist,  13,  435-438. 

1893  Test,  F.  C.  Bull.  U.S.  Fish  Commission,  H,  art.  I,  57-59. 

1895  Van  Denburgh,  J.  (see  Idaho). 

1932  Kellogg,  R.  Copeia,  no.  i,  p.  36. 

1932  Smith,  H.  M.  Copeia,  no.  2,  p.  100. 

1934  Donaldson,  L.  R.  Copeia,  no.  4,  p.  184. 

1942  Rodgers,  T.  L.,  and  W.  J.  Jellison.  Copeia,  no.  i,  pp.  10-11. 

1943  Slipp,  J.  W.,  and  G.  C.  Carl  (see  British  Columbia). 
Nebraska 

18573    Hallowell,  E.  Proc.  Acad.  Nat.  Sci.  Phila.  (Oct.  28,  1856),  8,  250-252. 
1875    Hayden,  F.  V.  Kept.  Engineer's  Dept.  U.S.  Army,  p.  105. 

Burt,  C.  E.,  and  M.  D.  Burt  (sec  Arkansas). 

Burt,  C.  E.,  and  M.  D.  Burt  (see  Arizona). 
1931     Burt,  C.  E.  (see  Kansas). 

1935  Burt,  C.  E.  (see  General  Worlds). 

1942    Hudson,  G.,  Univ.  Neb.  Conservation  and  Survey  Div.,  Neb.  Conserva- 
tion Bull.,  no.  24,  pp.  1-32,  105-124. 

1944  Marr,  J.  C.  (see  General  Worlds). 

1945  Loomis,  R.  Herpctologica,  27'8,  211-212. 
Nevada 

1878    Yarrow,  H.  C.,  and  H.  W.  Henshaw.  Geog.  Surveys  Territory  of  U.S. 

West  of  looth  Meridian,  App.  Ann.  Rept.,  pp.  206-210. 
1884    Cope,  E.  D.  Proc.  Acad.  Nat.  Sci.  Phila.  (1883),  35,  14,  18. 
1893    Stejneger,  L.  (see  California). 

1912  Taylor,  W.  P.  Univ.  Calif.  Pub.  Zool.,  7™,  340,  342-346. 

1913  Thompson,  H.  Proc.  Biol.  Soc.  Wash.,  26,  53-56. 

1915    Ruthven,  A.  G.,  and  H.  T.  Gaige.  Occ.  Papers  Mus.  Zool.  Univ.  Mich., 

no.  8,  pp.  1-34. 

1920    Snyder,  J.  O.  Copeia,  no.  86,  pp.  83-84. 
19213     Van  Denburgh,  J.,  and  J.  R.  Slevin.  Proc.  Calif.  Acad.  Sci.,  4th  ser.,  n2, 

27-30. 


BIBLIOGRAPHY  607 

1928  Slevin,  J.  R.  (see  British  Columbia). 

1932  Brues,  C.  T.  Proc.  Amer.  Acad.  Aits  Sciences,  67%  281-283. 

1932  Klauber,  L.  M.  (see  California). 

19333  Burt,  C.  E.  (see  General  Worlds). 

1936  Cowles,  R.  B.,  and  C.  M.  Bogert.  Herpetologica,  i33"42. 

1938  Linsdale,  J.  M.  Amer.  Midi.  Naturalist,  19*,  20-25. 

1940    Linsdale,  J.  M.  Proc.  Amer.  Acad.  Arts  Sciences,  73%  197-257. 

1942  Rivers,  I.  L.  Jour.  Ent.  &  ZooL  Pomona  College,  34%  52-56. 
New  Hampshire 

1899    Allen,  G.  M.  Proc.  Boston  Soc.  Nat.  Hist.,  29,  art.  3,  69-72. 

1904    Henshaw,  S.  (see  Connecticut). 

19180    Evermann,  B.  W.  Copeia,  no.  61,  pp.  81—83. 

1919    Speck,  F.  G.  Copeia,  no.  70,  pp.  46-48. 

1926  Babcock,  H.  L.  (see  Connecticut}. 

1939  Oliver,  J.,  and  J.  R.  Bailey.  In  Surv.  Rcpt.  N.H.  Pish  and  Game  Dept. 
Concord.  Vol.  IV,  pp.  195-217. 

1943  Aronson,  L.  R.  American  Midi.  Naturalist,  29%  242-244. 
New  Jersey 

1868     Abbott,  C.  C.  Geology  of  New  Jersey.  Newark.  App.  E,  pp.  804-805. 
1882    Abbott,  C.  C.  Amer.  Naturalist,  16,  707-711. 

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Abbott,  C.  C.  Amer.  Naturalist,  18,  1075-1080. 
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Peter,  J.  E.  Amer.  Naturalist,  23,  58-59. 
1890    Abbott,  C,  C.  Same,  24, 189. 

1890  Nelson,  J.  Geol.  Survey  N.J.,  Final  Rept.  State  Geol.,  2,  pt.  2.  Zoology, 
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1891  Cope,  E.  D.  Amer.  Natwalist,  25,  1017-1019. 

1904  Davis,  W.  T.  Same,  38,  893. 

1905  Davis,  W.  T.  Same,  39,  795,  796. 

1905  Ditmars,  R.  L.  (see  Connecticut). 

1906  Fowler,  H.  W.  Amer.  Naturalist,  40,  596. 

1906  Stone,  W.  (see  Delaware) . 

1907  Davis,  W.  T.  Amer.  Naturalist,  41,  49-50. 

1907  Fowler,  H.  W.  Ann.  Rept.  N.J.  State  Mus.  (1906),  pp.  1-408. 

1908  Davis,  W.  T.  Proc.  Staten  Is.  Assoc.  Arts  Sciences,  2,  pt.  II,  48-50. 

1908  Fowler,  H.  W.  Ann.  Rept.  N.J.  State  Mus.  (1907),  pp.  191-195- 

1909  Fowler,  H.  W.  Same  (1908),  pt.  Ill,  no.  2,  pp.  395-4°°- 

1910  Miller,  W.  DeW.,  and  J.  Chapin.  Science  (N.A.),  32S1«,  3i5-3r7- 
1914  Street,  J.  F.  Copeia,  no.  4,  p.  2. 

1916  Barbour,  T.  Copeia,  no.  26,  pp.  5-7. 

1916  Miller,  W.  D.  Copeia,  no.  34,  pp.  67,  68. 

1921  McAtee,  W.  L.  Copcia,  no.  96,  pp.  39,  40. 

1923  Noble,  G.  K.,  and  R.  C.  Noble.  Zoologica,  218,  414-455- 

1927  Brcder,  C.  M.,  Jr.,  R.  B.  Breder,  and  C.  Redmond.  Zoologica,  93,  201-229. 

Myers,  G.  S.  (see  Indiana). 


608  HANDBOOK  OF  FROGS  AND  TOADS 

1929  Myers,  G.  S.  Copeia,  no.  170,  p.  23. 

1930  Klots,  A.  B.  Copeia,  no.  173  (Oct.-Dcc.,  1929;  Jan.  16, 1930),  pp.  108-111. 

1931  Burt,  C.  E.  Jour.  Wash.  Acad.  Set.,  21°,  198-203. 

1936  Conant,  R.,  and  R.  M.  Bailey.  Occ.  Papers  Mus.  Zool.  Univ.  Mich.,  no. 
328,  pp.  1-5. 

1940    Moore,  J.  A.  (see  General  Worths). 
1943    Aronson,  L.  R.  Copeia,  no.  4,  pp.  246-249. 
1947    Blair,  A.  P.  Amer.  Mus.  Novitates,  no.  1343,  pp.  1-5. 
New  Mexico 

1854    Hallowell,  E.  Proc.  Acad.  Nat.  Sci.  Phila.  (Oct.  26, 1852),  6  (1852,  1853), 

181-182. 

1875    Yarrow,  H.  C.  (see  Arizona). 
1875    Coues,  E.  (see  Arizona). 

1884    Cope,  E.  D.  Proc.  Acad.  Nat.  Sci.  Phila.  (1883),  35,  10,  14,  15. 
1896    Cockerell,  T.  D.  A.  Amer.  Naturalist,  30,  327. 
1903    Stone,  W.,  and  J.  A.  G.  Rehn.  Proc.  Acad.  Nat.  Sci.  Phila. ,  55,  34. 
1907    Ruthven,  A.  G.  (see  Arizona). 
1913     Bailey,  V.  North  American  Fauna,  no.  35,  p.  35. 
1917    Ellis,  M,  M.  Copeia,  no.  43,  p.  39. 
ig24b    Van  Denburgh,  J.  Proc.  Caltf.  Acad.  Sci.,  4th  ser.,  i312, 189-191, 194-199. 

1932  Mosauer,  W.  Occ.  Papers  Mus.  Zool.  Univ.  Mich.,  no.  246,  pp.  1-5. 
19333    Burt,  C.  E.  (see  General  Wor\s). 

1933  Linsdale,  J.  M.  Copeia,  no.  4,  p.  222. 
1935    Burt,  C.  E.  (see  General  Wor\s). 

1937  Little,  E.  L.,  Jr.,  and  J.  G.  Keller.  Copeia,  no.  4,  pp.  216-222. 

1938  Taylor,  E.  H.  (see  Arizona). 

I94ib    Bragg,  A.  N.  Great  Basin  Naturalist,  2%  109-1 17. 
i94ic    Bragg,  A.  N.  Wasmann  Collector,  4%  92-94. 
1946    Koster,  W.  J.  Copeia,  no.  3,  p.  173. 
New  York 

1842    DeKay,  J.  E.  Zoology  of  New  Yor\,  or  New  Yorl(  Fauna.  Pt.  Ill,  "Rep- 
tiles and  Amphibia,"  pp.  59-72.  Pt.  IV,  plates  19-22. 

1851  Gebbard,  J.,  Jr.  N.Y.  State  Cabinet  Nat.  Hist.,  p.  23. 

1852  Hough,  F.  B.  Same,  fifth  ann.  rept.,  p.  19. 

1852  Hough,  F.  B.  Same,  p.  24. 

1853  Gebbard,  J.,  Jr.  Same,  p.  23. 

1878  Monks,  S.  P.  Amer.  Naturalist,  12,  695. 

1879  Gilbert,  E.  S.  Science  News,  p.  336. 

1882  Bicknell,  E.  P.  Trans.  Linn.  Soc.  N.Y.,  i,  124. 

1898  Davis,  W.  T.  Proc.  Nat.  Sci.  Assoc.  Staten  Is.,  f,  4. 

1898  Mearns,  E.  A.  Bull.  Amer.  Mus.  Nat.  Hist.,  no.  10,  art.  XVI,  pp.  324-326. 

1898  Sherwood,  W.  L.  (see  Connecticut). 

1899  Mearns,  E.  A.  Proc.  U.S.  Nat.  Mus.,  21, 345.  (Read  Nov.  4,  1898.) 

1902  Paulmier,  F.  C.  N.Y.  State  Mus.  Bull.,  no.  51,  pp.  403, 414. 

1903  Britcher,  H.  W.  Proc.  Onondaga  Acad.  Sci.,  i,  120-122. 

1904  Gage,  S.  H.  Cornell  Nature-Study  Leaflet,  no.  16,  pp.  185-205;  Teachers' 
Leaflet,  no.  9  (May,  1897). 


BIBLIOGRAPHY  609 

1905  Ditmars,  R.  L.  (see  Connecticut). 

1908  Wright,  A.  H.,  and  A.  A.  Allen.  Amer.  Naturalist,  42,  39-42. 

1910  Miller,  W.  DeW.,  and  }.  Chapin  (see  New  Jersey). 

1912  Davis,  W.  T.  Proc.  Staten  Is.  Assoc.  Arts  Sciences  (Oct.  igog-May  1911), 

pp.  66-67.  Read  Apr.  16, 1910. 

1914  Overton,  F.  Mus.  Brooklyn  Inst.  Arts  Sciences,  Science  Bull.,  28,  23-40. 

1914  Wright,  A.  H.  Carnegie  Inst.  Pub.,  no.  197,  pp.  1-98. 

1915  Overton,  F.  Copeia,  no.  20,  p.  17;  no.  24,  pp.  52-53. 

1917  Nichols,  J.  T.  Copeia,  no.  45,  pp.  59-60. 

1918  Evermann,  B.  W.  Copeia,  no.  56,  pp.  48-51. 

1918  Latham,  R.  Copeia,  no.  62,  p.  88. 

1919  Wright,  A.  H.,  and  J.  Moesel,  Copeia,  no.  74,  pp.  81-83. 

1920  Fisher,  G.  C.  Copeia,  no.  85,  pp.  76-77. 
1923  Bishop,  S.  C.  Copeia,  no.  120,  pp.  83-84. 

1923  Green,  H.  T.  Copeia,  no.  122,  p.  99. 

1924  Frost,  S.  W.  Jour.  N.Y.  Ent.  Soc.,  32,  173-185. 

1927  Bishop,  S.  C.,  and  W.  P.  Alexander.  N.Y.  State  Museum  Handbool^,  no.  3. 

Albany.  Pp.  60-76. 

1927  Heller,  J.  A.  Copeia,  no.  165,  p.  1 16. 

1927!?  Myers,  G.  S.  (see  Indiana). 

1828  Weber,  J.  A.  Copeia,  no.  169,  pp.  106,  108-110. 

1930  Hamilton,  W.  J.,  Jr.  Copeia,  no.  2,  p.  45. 

1930  Myers,  G.  S.  (see  New  Jersey). 

1931  Burt,  C.  E.  (see  New  Jersey). 

1932  Frost,  S.  W.  (see  Pennsylvania). 

1934  Hamilton,  W.  J.,  Jr.  Copeia,  no.  2,  pp.  88-90. 

1934  Maynard,  E.  A.  Copeia,  no.  4,  pp.  174-177. 

1935  Frost,  S.  R.  (see  Pennsylvania). 

1936  Kauflfeld,  C.  F.  Herpetologica,  i,  IT. 
1939  Janes,  R.  G.  Copeia,  no.  3,  pp.  134-140. 

1939  Moore,  J.  A.  Ecology,  2o4,  459-478. 

1940  Raney,  E.  C.,  and  W.  M.  Ingram.  Bull.  Ecol.  Soc.  Amer.,  2i4, 30. 
1940  Raney,  E.  C.  Amer.  Midi.  Naturalist,  23 J,  733-745- 

1940  Moore,  J.  A.  (see  General  Worlds). 

1941  Moore,  J.  A.  (see  General  Worlds). 
1941  Pratt,  J.  Copeia,  no.  4,  p.  264. 

1941  Raney,  E.  C.,  and  W.  M.  Ingram.  Amer.  Midi.  Naturalist,  261,  201-206. 

1941  Raney,  E.  C.,  and  W.  M.  Ingram  (see  General  Worfo). 
1942 '  Gilbert,  P.  W.  Copeia,  no.  3,  p.  177. 

1942  Vogel,  H.  H.,  Jr.  Proc.  Ind.  Acad.  Sci.  (1941),  p.  266. 

1943  Aronson,  L.  R.  Amer.  Mus.  Novitates,  no.  1224,  pp.  1-6. 
1943  Clausen,  R.  T.  Amer.  Midi.  Naturalist,  29*,  360-364. 

1943  Ingram,  W.  M.,  and  E.  C.  Raney.  Same,  29%  239-241. 

1944  Aronson,  L.  R.  Amer.  Mus.  Novitates,  no.  1250,  pp.  1-15. 

1944  Moore,  C.  B.  Science  Digest,  15%  75,  76;  also,  Nat.  History,  50  (1942), 
191—192. 

1945  Eaton,  T.  H.,  Jr.  Copeia,  no.  2,  p.  115. 


6io  HANDBOOK  OF  FROGS  AND  TOADS 

1947  Blair,  A.  P.  (see  New  Jersey). 

1947  Evans,  H.  E.  Herpetologica,  4,  19-21. 

1947  Raney,  E.  C.,  and  E.  A.  Lachner.  Copeia,  1947,  pp.  113-116. 
North  Carolina 

1870  Cope,  E.  D.  Amer.  Naturalist,  4, 397. 

1878  Coues,  E.,  and  H.  C.  Yarrow.  Proc.  Acad.  Nat.  Sci.  Phtta.,  30,  28. 

1879  Humphreys,  J.  T.  Science  News,  p.  304. 
1891  Ryder,  J.  A.  Amer.  Naturalist,  25,  838-840. 
1896  Brimley,  C.  S.  Same,  30,  501. 

1907-1927  Brimley,  C.  S.  Jour.  Elisha  Mitchell  Sci.  Soc.,  2$*,  157-160;  2d 
ed.,  1926,  421""2,  80-83. 

1908  Brimley,  C.  S.,  and  F.  Sherman,  Jr.  Same,  24,  14-22. 

1909  Brimley,  C.  S.  Proc.  Biol.  Soc.  Wash.,  22, 129-131, 133. 

1910  Brimley,  C.  S.  (see  Alabama). 

1917  Dunn,  E.  R.  Bull.  Amer.  Mus.  Nat.  Hist.,  37,  593-597,  620-623. 

1920  Dunn,  E.  R.  (see  Alabama). 

1921  Metcalf,  Z.  P.  Copeia,  no.  100,  pp.  81-82. 

1922  Brimley,  C.  S.  Copeia t  no.  107,  p.  48. 

1923  Breder,  C.  M.,  Jr.,  and  R.  B.  Breder.  Zoologica,  4,  3-9,  18-20. 

1923  Brimley,  C.  S.  Copeia,  no.  114,  p.  4. 

1924  Myers,  G.  S.  Copeia,  no.  131,  pp.  59-60. 

1925  Brimley,  C.  S.,  and  W.  B.  Mabie.  Copeia,  no.  139,  p.  15. 
19273    Brimley,  C.  S.  Copeia,  no.  162,  pp.  11-12. 

i927b    Myers,  G.  S.  (see  Indiana). 

1928    Andrews,  E.  A.  Science,  new  ser.,  67, 269-270. 

1928     Bishop,  S.  C.  Jour.  Elisha  Mitchell  Sci.  Soc.,  43,  167. 

1933     Brandt,  B.  B.  Copeia,  no.  i,  p.  39. 

1933    Brandt,  B.  B.,  and  C.  F.  Walker.  Occ.  Papers  Mus.  Zool.  Univ.  Mich., 

no.  272,  pp.  1-7. 

1935     Harper,  F.  (see  General  Worlds). 
19363    Brandt,  B.  B.  Copeia,  no.  4,  pp.  215-223. 
i936b    Brandt,  B.  B.  Ecol.  Monog.  (Durham,  N.C.),  6,  491-532. 
1939    King,  W.  (see  Tennessee). 

1941  Gray,  I.  E.  Amer.  Midi.  Naturalist,  25',  652-658. 

1940-1941  Brimley,  C.  S.  Carolina  Tips,  3  (April,  1940),  10-11;  (June,  1940), 
14,  15;  (August,  1940),  18-19;  (Sept.,  1940),  22-23;  (Nov.,  1940),  26- 
27;  (Jan.,  1941),  2-3. 

1942  Engels,  W.  L.  Amer.  Midi.  Naturalist,  28,  293-294. 
North  Dakota 

1874  Allen,  J.  A.  (see  Montana). 

1878  Coues,  E.,  and  H.  C.  Yarrow  (see  Montana). 

1887  Ballou,  W.  H.  Amer.  Naturalist,  21,  388. 

1944  Telford,  H.  S.  N.D.  Ag.  Expt.  Stat.  Bimonthly  Bull.,  64,  33-35. 

1944  Telford,  H.  S.  Same,  6%  35-37. 
Ohio 

1838  Kirtland,  J.  P.,  "Rept.  on  Zoology  of  Ohio."  In  W.  W.  Mather,  Second 
Ann.  Rept.  Geol.  Surv.  Ohio.  Columbus.  P.  168. 


BIBLIOGRAPHY  611 

1882    Smith,  W.  H.  Reft.  Geol.  Survey  Ohio,  4,  pt.  I,  2ool.  sect.  Ill,  pp.  701-713, 

733>  734- 
1887    James,  J.  F.  Jour.  Cincinnati  Soc.  Nat.  Hist.,  ic1, 35-36. 

1891     Wilcox,  E.  V.  Otterbein  (Ohio)  Aegis. 

i8g5a    Kirsch,  P.  H.  Bull.  U.S.  Fish  Commission  (1894),  *4»  art-  2O>  333- 

1901     Morse,  M.  Ohio  Naturalist,  i7,  114-115. 

1903  Morse,  M.  Same,  3'%  360,  361. 

1904  Morse,  M.  Proc.  Ohio  State  Acad.  Sci.,  4,  pt.  3,  Special  papers  no.  9, 
102,  116-122. 

1932  Walker,  C.  F.  Ohio  Jour.  Sci.,  32%  379-384. 

1933  Walker,  C.  F.  Copeia,  no.  4,  p.  224. 

J937    Conant,  R.  P.,  and  W.  M.  Clay.  Occ.  Papers  Mus.  Zool.  Univ.  Mich.,  no. 
346,  pp.  1-9. 

1945  Gier,  H.  T.  Copeia,  no.  i,  p.  50. 

1946  Walker,  C.  F.  Ohio  State  Arch.  Hist.  Soc.t  Ohio  State  Mus.  Bull,  i3, 
1-109. 

1947  Wood,  J.  T.,  and  W.  E.  Duellman.  Heipetologica,  4,  3-6. 
Oklahoma 

1894  Cragin,  F.  W.  (see  Kansas). 

1894  Cope,  E.  D.  Pi oc.  Acad.  Nat.  Sci.  Phila.  ( 1893),  45»  386- 

1903  Stone,  W.  (see  Arkansas). 

1919  Schmidt,  K.  P.  Copeia,  no.  73,  p.  71. 

1924  Ortenburger,  A.  I.  Proc.  Otya.  Acad.  Sci.,  4,  19-20. 

1925  Force,  E.  R.  Copeia,  no.  141,  p.  25. 

1926  Force,  E.  R.  Proc.  O%la.  Acad.  Sci.f  5,  80-82.  Univ.  OJ(la.  Bull.,  new  ser., 
no.  330,  Univ.  Studies  no.  22. 

iQ26a    Ortenburger,  A.  I.  Copeia,  no.  155,  pp.  137-138. 
i926b    Ortenburger,  A.  I.  Copeia,  no.  156,  p.  145. 

Ortenburger,  A.  I.  Proc.  O\la.  Acad.  Sci.,  6,  pt.  I,  89-93.  Univ.  Of(la. 
Bull.,  new  ser.,  no.  348. 

Ortenburger,  A.  I.  Copeia,  no.  163,  pp.  46-47. 

1928    Force,  E.  R.  Proc.  Otya.  Acad.  Set.,  art.  XVII,  pp.  78,  79.  Univ.  Oltfa.  Bull., 
new  ser.,  no.  410,  Univ.  Studies  no.  50. 
Burt,  C.  E.,  and  M.  D.  Burt  (see  Atfonsas). 
Ortenburger,  A.  I.  Copeia,  no.  170,  pp.  8-10. 
Ortenburger,  A.  I.  Copeia,  no.  170,  pp.  26-27. 
1930    Force,  E.  R.  Copeia,  no.  2,  pp.  25-27,  38,  39. 

1930    Ortenburger,  A.  I.,  and  Beryl  Freeman.  Pub.  Univ.  Ol(la.,  2,  Biol.  Survey, 
no.  4, 176-178. 

1930  Ortenburger,  A.  I.  Copeia,  no.  173  (Oct.-Dec.  1929;  Jan.  16,  1930),  p.  94. 

193 1  Burt,  C.  E.  (  see  Kansas) . 

1933  Smith,  H.  M.  (see  Kansas). 

1934  Smith,  H.  M.,  and  A.  B.  Leonard.  Amer.  Midi.  Naturalist,  15,  190-196. 

1935  Burt,  C.  E.  (see  General  Wor\s). 

1936  Blair,  W.  F.  Copeia,  no.  2,  p.  115. 
1936  Bragg,  A.  N.  Copeia,  no.  i,  pp.  14-20. 
J937  Bfagg»  A.  N.  Copeia,  no.  4,  pp.  227-228. 


612  HANDBOOK  OF  FROGS  AND  TOADS 

19373    Bragg,  A.  N.  Amer.  Midi.  Naturalist,  i82,  273-284. 

1937    Moore,  G.  A.  Copeia,  no.  4,  pp.  225-226. 

1937    Trowbridge,  A.  H.  Copeia,  no.  i,  pp.  71-72. 

1937    Trowbridge,  A.  H.  Amer.  Midi.  Naturalist,  i82,  284-303. 

1937    Trowbridge,  A.  H.  and  M.  S.  Trowbridge.  Amer.  Naturalist,  71™,  460- 

480. 

1939    Bragg,  A.  N.  Proc.  Ot(la.  Acad.  Set.  19,  41-42. 
19393    Bragg,  A.  N.  Copeia,  no.  3,  p.  173. 

1939  Jones,  R.  W.,  and  G.  E.  Derrick.  Proc.  Oltfa.  Acad.  Sci.  19, 39. 
i94oa    Bragg,  A.  N.  Amer.  Naturalist,  74,  322-349,  424-438. 
i94ob    Bragg,  A.  N.  Wasmann  Collector,  4,  6-16. 

19400    Bragg,  A.  N.  Proc.  O\la.  Acad.  Sci.,  20,  71-74. 

I94od    Bragg,  A.  N.  Same,  pp.  75-76. 

I94oe    Bragg,  A.  N.  Observations  on  the  Ecology  of  Natural  History  of  Anura 

H  and  111.  Amer.  Midi.  Naturalist,  242,  306-335.  (Observation  II,  pp. 

301-321;  Observation  III,  pp.  322-335.) 

1940  Bragg,  A.  N.,  and  R.  E.  Kuntz.  Proc.  Otya.  Acad.  Sci.,  20,  77. 

1941  Bragg,  A.  N.  Key  to  the  Toads  (Bufo)  of  Oklahoma.  Same  (1940),  21, 
17-18. 

19413    Bragg,  A.  N.  Copeia,  no.  i,  pp.  51-52. 

i94ib    Bragg,  A.  N.  (see  New  Mexico). 

I94id    Bragg,  A.  N.  Turtox  News,  19%  10-12,  (Observation  VIII.) 

1941    Smith,  C.  C.,  and  A.  N.  Bragg  (see  General  Worlds). 

19413    Smith,  C.  C.,  and  A.  N.  Bragg  (see  General  Worlds). 

1941  Trowbridge,  M.  S.  Trans.  Amer.  Micros.  Soc.,  6o4,  508-526. 

1942  Bragg,  A.  N.  Proc.  Ofya.  Acad.  Sci.,  22,  16-17. 
19423    Bragg,  A.  N.  Proc.  Of(la.  Acad.  Sci.,  22,  18. 
19420    Bragg,  A.  N.  Turtox  News,  20%  12,  13. 
i942d    Bragg,  A.  N.  Same,  2O11,  154. 

19426    Bragg,  A.  N.  Science,  new  ser.,  95,  194-195. 
i942f    Bragg,  A.  N.  Anat.  Rec.,  84*,  506. 
i942g    Bragg,  A.  N.  Same,  84%  507. 
i942h    Bragg,  A.  N.  Wasmann  Collector,  5'-,  47-60. 
1942!    Bragg,  A.  N.  Proc.  O^la.  Acad.  Sci.,  22,  73-74.  (Observation  XI.) 
1942    Bragg,  A.  N.,  and  C.  C.  Smith.  Great  Basin  Naturalist,  22,  33-50.  (Obser- 
vation IX.) 
1942    Moore,  G.  A.,  and  C.  C.  Rigney.  Proc.  OJ{la.  Acad.  Sci.  (1941),  22,  77-80. 

1942  Trowbridge,  M.  S.  Trans.  Amer.  Micros.  Soc.,  61%  66-83. 

1943  Blair,  A.  P.  Amer.  Naturalist,  77%  563-568. 

1943  Bragg,  A.  Great  Basin  Naturalist,  43'4,  62-80.  (Observation  XV.) 

19433  Bragg,  A.  N.  Proc.  Ofa.  Acad.  Set.  (1942),  23,  37-39. 

i943b  Bragg,  A.  N.  Same,  pp.  39-40. 

19430  Bragg,  A.  N.  Wasmann  Collector,  5*, 129-140.  (Observation  XVI.) 

1943  Bragg,  A.  N.,  and  C.  C.  Smith.  Turtox  News,  21 9,  107. 

1944  Bragg,  A.  N.  Copeia,  no.  4,  pp.  230-241.  (Observation  XIII.) 
19443  Br3gg,  A.  N.  Proc.  O\la.  Acad.  Sci.  (1934),  24, 13-14. 


BIBLIOGRAPHY  613 

I944b    Bragg,  A.  N.  Amer.  Naturalist,  78,  517-533;  79>  52-72.  (Observation 
XII.) 

1944  Marr,  J.  C.  (see  General  Worths). 

1945  Bragg,  A.  N.  Proc.  Oltfa.  Acad.  Sci.  (1944),  25,  27. 

1945  Bragg,  A.  N.  Wasmann  Collector,  6**-  *,  68^-78. 

1946  Bragg,  A.  N.  Proc.  Otya.  Acad.  Sci.,  26,  16. 
i946a    Bragg,  A.  N.  Proc.  Otya.  Acad.  Sci.,  26,  19. 
i946b    Bragg,  A.  N.  Herpetologica,  3,  89-97. 

1946  Dundee,  H.,  and  A.  N.  Bragg.  Proc.  Ol(la.  Acad.  Sci.t  26, 18. 
1945    Smith,  H.  M.  Univ.  Kans.  Pub.  Mus.  Nat.  Htst.,  i3,  88-89. 

1947  Blair,  A.  P.  (see  New  Jersey). 
Oregon 

1854  Baird>  s-  F->  and  c-  CJirard.  Proc.  Acad.  Nat.  Sci.  Phtla.  (Oct.  26,  1842), 

6,(i852, 1853),  174-175. 

1854  Hallowell,  E.  Same,  p.  183. 

1884  Cope,  E.  D.  Same  (1883),  35, 19, 20,  23. 

1895  Van  Denburgh,  J.  (see  Idaho). 

1899  Washburn,  F.  L.  Amer.  Naturalist,  33™°,  139-141. 
I9i2a    Van  Denburgh,  J.  (see  Idaho). 

1917  Camp,  C.  L.  Copeia,  no.  40,  pp.  13,  14. 

1922  Van  Winkle,  K.  (see  Washington). 

1928  Brues,  C.  T.  Proc.  Amer.  Acad.  Arts  Sciences,  63*,  205-206. 

1928  Slevin,  J.  R.  (see  British  Columbia). 

1934  Brooking,  W.  J.  Copeia,  no.  2,  pp.  93,  94. 

1936  Fitch,  H.  S.  Amer.  Midi.  Naturalist,  17',  634-652. 

1936  Jewett,  S.  G.,  Jr.  Copeia,  no.  i,  pp.  71,  72. 

1938  Fitch,  H.  S.  Copeia,  no.  3,  p.  148. 

1939  Gordon,  K.  Oregon  State  Coll.  Monog.,  no.  i,  pp.  1-82. 

1939    Graf,  W.,  Stanley  G.  Jewett,  Jr.,  and  Kenneth  Gordon.  Copeia,  no.  2, 
pp.  101-104. 

1942  Slater,  J.  R.  (see  Washington). 

1943  Evenden,  F.  G.,  Jr.  Copeia,  no.  4,  pp.  251-252. 
1945     Schonberger,  C.  F.  Copeia,  no.  2,  pp.  120-121. 

Pennsylvania 

1843  Haldeman,  S.  S.  In  Trego,  Geography  of  Pennsylvania.  Philadelphia.  P. 
78. 

1844  Haldeman,  S.  S.  In  Rupp,  History  of  Lancaster  County.  Lancaster,  Pa. 
Appendix. 

1857  Hallowell,  E.  Proc.  Acad.  Nat.  Hist.  Phila.  (1856),  8, 141-142. 

1869  Stauffer,  J.  In  Mombert,  History  of  Lancaster  County.  Lancaster,  Pa. 

P.  576. 

1879  Rath  von,  S.  S.  Science  News,  p.  368. 

1903  Reese,  A.  M.  Science,  new  sen,  i7425. 

1906  Stone,  W.  (see  Delaware). 

1908  Palmer,  T.  C.  Proc.  Delaware  Co.  (Pa.)  Inst.  Sci.,  41,  12-22. 

1913  Surface,  H.  A.  Pa.  Dept.  Ag.,  Div.  Zool.,  Bimonthly  Zool.  Bull.,  3, 66-152. 


614  HANDBOOK  OF  FROGS  AND  TOADS 

1914  Palmer,  T.  C.  Proc.  Delaware  Co.  (Pa.)  Inst.  Sci.,  7, 15-17. 

1915  Dunn,  E.  R.  Copeia,  no.  16,  p.  3. 

1915  Fowler,  H.  W.  Proc.  Delaware  Co.  (Pa.)  Inst.  Sci.,  f,  41,  42. 

1915  Keim,  T.  D.  Copeia,  no.  24,  pp.  51-52. 

1917  Fowler,  H.  W.  Copeia ,  no.  40,  pp.  14,  15, 

1917  Mattern,  E.  S.,  and  W.  I.  Mattern.  Copeia,  no.  46,  p.  65. 

191 8b  Evermann,  B.  W.  Copeia,  no.  58,  p.  67. 

1924  Frost,  S.  W.  (see  New  Yorty. 

1928  Netting,  M.  G.  The  Cardinal,  2%  2. 

1932  Frost,  S.  W.  Amer.  Naturalist,  66,  530-540. 

1933  Burger,  J.  W.  Copeia,  no.  2,  pp.  94. 

1933  Netting,  M.  G.  Proc.  Pa.  Acad.  Sci.,  7,  i,  7-11. 

1935  Frost,  S.  R.  Copeia,  no.  i,  pp.  15-18. 

19363    Netting,  M.  G.  Proc.  Pa.  Acad.  Set.,  10,  26-28. 

i936b    Netting,  M.  G.  Carnegie  Mus.  Pa.  Her  fetology  Leaflet,  no.  i. 

1939  Mohr,  C.  E.  Proc.  Pa.  Acad.  Sci.,  13,  76-78. 

1939  Pawling,  R.  O.  Herpetologtca,  i,  165-169. 

1940  Mohr,  C.  E.  Frontiers,  4%  142-146. 

1940  Yoder,  H.  D.  Proc.  Pa.  Acad.  Sci.,  14,  90-92. 

1941  Frey burger,  W.  A.,  Jr.  Same,  15,  180-183. 

1941  Knepp,  T.  H.  Same,  p.  164. 

1942  Conant,  R.  Amer.  Midi.  Naturalist,  27%  161-163. 

1943  Baldauf,  R.  J.  Mengel  Nat.  Hist.  Soc.  Leaflet,  no.  i,  pp.  1-8. 

1945  Smith,  A.  G.  Proc.  Pa.  Acad.  Sci.f  19,  77. 
Rhode  Island 

1884-1886     Bumpus,  H.  C.  Random  Notes  of  Natural  History,  i10,  4,  5;  3% 

52. 

1904  Henshaw,  S.  (see  Connecticut). 

1905  Drowne,  F.  P.  Roger  Williams  Mar^  Mus.  Monogr.,  no.  15,  pp.  1-24. 
1921     Babcock,  H.  L.  (see  Connecticut). 

1926    Babcock,  H.  L.  (see  Connecticut). 
South  Carolina 

1920    Deckert,  R.  F.  Copeia,  no.  80,  p.  26. 
i924b    Schmidt,  K.  P.  Copeia,  no.  132,  p.  68. 

1924     McManus,  I.  The  Anura  of  the  Columbia,  S.C.  Region.  M.  A.  Thesis, 
Columbia. 

Pickens,  A.  L.  Copeia,  no.  162,  pp.  25-26. 
Pickens,  A.  L.  Copeia,  no.  165,  pp.  106-110. 

1929  Corrington,  J.  D.  Copeia,  July-Sept.,  pp.  58-67. 
1935     Harper,  F.  (see  General  Worlds). 

1937    Chamberlain,  E.  B.  Copeia,  no.  2,  p.  142. 

1939  Chamberlain,  E.  B.  Charleston  Mus.  Leaflet,  no.  12,  pp.  1-38. 

1940  Jopson,  H.  G.  M.  Herpetologica,  2,  pt.  2,  39-43. 
1940    Stejneger,  L.  (see  Florida). 

1946  Obrecht,  C.  B.  Copeia,  no.  2,  pp.  71-72. 

1947  Blair,  A.  P.  (see  New  Jersey). 
i948b    Neill,  W.  T.  (see  Georgia). 


BIBLIOGRAPHY  615 

South  Dakota 

1874    Allen,  J.  A.  (see  Montana). 

1874  Coues,  E.  (see  Montana). 

1875  Hayden,  R  V.  (see  Nebraska). 

1908    Reagen,  A.  B.  Zoologischer  Anxeiger,  32,  Band  Nr.  I,  31;  also  in  Rept. 

State  GeoL,  S.D.  GcoL  Survey  Bull.,  no.  4,  p.  164. 
1914  Visher,  S.  S.  S.D.  GeoL  Survey  Bull.,  no.  6,  p.  93. 
1923  Over,  W.  H.  5.D.  GeoL.Nat.  Hist.  Survey  Bull.,  12,  Bull.  Univ.  S.D.  ser. 

XXIII,  no.  10,  pp.  11-15. 
19293     Burt,  C.  E.,  and  M.  D.  Burt  (see  Arkansas). 

1935  Burt,  C.  E.  (see  General  Wor^s). 

1943  Larsen,  N.  P.  Jour.  Econ.  Ent.  63,  no.  3,  480. 
Tennessee 

1844    Troost,  G.  Seventh  GeoL  Rept.  Tenn.,  p.  40. 

1896    Rhoads,  S.  N.  Proc.  Acad.  Nat.  ScL  Phtla.,  47  (Nov.  15,  1895),  376-383, 

395-399,  4°5>  4<>7- 
1920    Dunn,  E.  R.  (see  Alabama). 

1922  Blanchard,  F.  N.  Occ.  Papers  Mus.  ZooL  Untv.  Mich.,  no.  117,  pp.  4-6. 

1927  Dunn,  E.  R.  Copeia,  no.  162,  p.  19. 

1934  Necker,  W.  Bull.  Chicago  Acad.  ScLt  5%  1-4. 

1936  Bailey,  J.  R.  Copeia,  no.  2,  p.  115. 

1937  Cagle,  F.  R.  Jour.  Tenn.  Acad.  Sci.,  I22,  179-185. 
1937  Endsley,  J.  R.  Copeia,  no.  i,  p.  70. 

19373    Parker,  M.  V.  Jour.  Tenn.  Acad.  Sci.f  12,  169-178. 

i937b    Parker,  M.  V.  Same,  I21,  60-86. 

1939    Cahn,  A.  R.  Copeia,  no.  r,  pp.  52-53. 

1939    King,  W.  Amer.  Midi.  Naturalist,  23,  531-582. 

1939  Parker,  Malcolm  V.  Jour.  Tenn.  Acad.  ScL,  I41,  72,  73,  76-81. 

1940  Shoup,  C.  S.,  and  J.  H.  Peyton.  Jour.  Tenn.  Acad.  ScL,  15,  114. 

1941  Gentry,  G.  Tenn.  Dept.  Conservation,  Div.  Fish  and  Game,  Misc.  Pub., 
no.  4,  pp.  329-331. 

1944  King,  W.  Copeia,  no.  4,  p.  255. 
Texas 

1854    Baird,  S.  F.,  and  C.  Girard.  Proc.  Acad.  Nat.  ScL  Phila.  (1852),  6  (1852, 

1853)*  173- 

1857!)    Hallowell,  E.  Same  (1856),  8,  307-309. 
1859    Baird,  S.  F.  "Reptiles  of  the  Boundary."  In  W.  H.  Emory,  U.S.  and  Mex. 

Boundary  Survey.  Vol.  II  (1858),  25-29. 
1859    Baird,  S.  F.  (see  Arkansas). 

1865    Cope,  E.  D.  Proc.  Acad.  Nat.  ScL  Phila.,  17, 194,  195, 197. 
1878    Cope,  E.  D.  Amer.  Naturalist,  12,  186;  i24,  252,  253. 
i88oa    Cope,  E.  D.  Bull.  U.S.  Nat.  Mus.,  no.  17,  pp.  8,  24-29,  42-47. 
i88ob    Cope,  E,  D.  Proc.  Amer.  Phil.  Soc.,  18  (June  20,  1879),  p.  263. 

1888  Carman,  S.  Bull.  Essex  Inst.,  19  (1887),  134,  138. 

1889  Cope,  E.  D.  Proc.  U.S.  Nat.  Mus.  (1888),  u,  317-318,  395~396- 
1893    Cope,  E.  D.  Proc.  Acad.  Nat.  ScL  Phila.  (1892),  44,  333,  337. 
1893    Cope,  E.  D,  Amer,  Naturalist,  no.  314,  pp.  155-156. 


616  HANDBOOK  OF  FROGS  AND  TOADS 


1894    Test>  F-  c-  Bul1-  u-s-  FisA  Commission,  12  (1892,  1894),  122. 
1899    Mocquard,  F.  (sec  General  Worlds). 

1902  Strecker,  J.  K.,  Jr.  Trans.  Tex.  Acad.  Sci.  (1901),  4%  pt.  II,  6-7. 

1903  Stone,  W.  (see  Arkansas). 

1905    Gadow,  H.  Proc.  Zool.  Soc.  London,  2,  193-194,  205-208,  230-231. 
1907    Mearns,  E.  A.  Bull.  U.S.  Nat.  Mus.,  no.  56,  pt.  I,  p.  81. 
igoSa    Strecker,  J.  K.,  Jr.  Proc.  Biol.  Soc.  Wash.,  21,  80-83. 
I9o8b    Strecker,  J.  K.,  Jr.  Same,  pp.  51-52. 

Strecker,  J.  K.,  Jr.  Same,  pp.  56-61. 

Strecker,  J.  K.,  Jr.  Same,  p.  88. 
I9o8e    Strecker,  J.  K.,  Jr.  Same,  pp.  199-206. 
1909    Strecker,  J.  K.,  Jr.  Baylor  Univ.  Bull.,  12%  9,  15. 
19093    Strecker,  J.  K.,  Jr.  Proc.  Biol.  Soc.  Wash.,  22,  115-120. 
19103    Strecker,  J.  K.,  Jr.  Same,  23,  115-122. 
i9iob    Strecker,  J.  K.,  Jr.  Baylor  Univ.  Bull.,  13*-  n,  1-4,  17-21. 
i9ioc    Strecker,  J.  K.,  Jr.  Trans.  Acad.  Sci.  St.  Louis,  19*,  73-82. 
1915    Stejneger,  L,  Proc.  Biol.  Soc.  Wash.,  28,  131-132. 
1915     Strecker,  J.  K.,  Jr.  Baylor  Univ.  Bull.,  18*,  45-54,  61-82. 

1919  Pope,  P.  H.  Copeia,  no.  76,  pp.  93-98. 

1920  Schmidt,  K.  P.  (see  Louisiana). 

1922    Strecker,  J.  K.,  Jr.  Bull.  Sci.  Soc.  San  Antonio,  no.  4,  pp.  1-5,  9-16,  45. 

19263    Strecker,  J.  K.,  Jr.  Cont.  Baylor  Univ.  Mus.,  no.  2,  pp.  i,  2. 

i926b    Strecker,  J.  K.,  Jr.  Same,  no.  3,  p.  3. 

i926c    Strecker,  J.  K.,  Jr.  Same,  no.  6,  pp.  i,  8,  9. 

i926d    Strecker,  J.  K.,  Jr.  Same,  no.  7,  pp.  3-11. 

19260    Strecker,  J.  K.,  Jr.  Same,  no.  7,  pp.  8-n. 

ig26f    Strecker,  J.  K.,  Jr.  Same,  no.  8,  pp.  7-12. 

19273    Strecker,  J.  K.,  Jr.  Same,  no.  10,  pp.  13,  14. 

i927b    Strecker,  J.  K.,  Jr.  Copeia,  no.  162,  p.  8. 

1927  Strecker,  J.  K.,  Jr.,  and  W.  J.  Williams.  Cont.  Baylor  Univ.  Mus.,  no.  12, 
pp.  1-8,  13,  1  6. 

1928  Strecker,  J.  K.,  Jr.,  and  W.  J.  Williams.  Same,  no.  17,  pp.  3-19. 
i928b    Strecker,  J.  K.,  Jr.  Same,  no.  15,  pp.  3-5,  7. 

i928c    Strecker,  J.  K.,  Jr.  Same,  no.  16,  pp.  1-21,  3-6,  9-10. 
19293    Burt,  C.  E.,  and  M.  D.  Burt  (see  Arkansas). 
i929b    Burt,  C.  E.,  and  M.  D.  Burt  (see  Arizona). 

1929  Strecker,  J.  K.,  Jr.  Cont.  Baylor  Univ.  Mus.,  no.  19,  pp.  1-15. 

1930  Strecker,  J.  K.,  Jr.  Same,  no.  23,  pp.  2-4,  6-8. 

1931  Gaige,  H.  T.  Copeia,  no.  2,  p.  63. 

1932  Burt,  C.  E.  Copeia,  no.  3,  p.  158. 
19323    Burt,  C.  E.  Bio-Log,  21,  i,  2. 
1932    Mosauer,  W.  (see  New  Mexico). 

1932    Taylor,  E.  H.  Univ.  Kans.  Sci.  Bull.,  no.  u,  pp.  243-245. 

1932  Taylor,  E.  H.,  and  J.  S.  Wright.  Same,  no.  12,  pp.  247-249. 

1933  Martin,  F.  Aquar.  (Berlin),  pp.  92-93. 
1933    Smith,  H.  M.  Copeia,  no.  4,  p.  217. 

1933    Strecker,  J.  K.,  Jr.  Copeia,  no.  2,  pp.  77-79. 


BIBLIOGRAPHY  617 

1934  piatt>  J-  Amer.  Midi.  Naturalist,  15*,  89-91. 

1935  Strecker,  J.  K.,  Jr.,  and  J.  E.  Johnson.  Baylor  Univ.  Bull.,  no.  38,  3,  pp. 
17-23. 

1935  Williams,  W.  J,  Baylor  Bulletin,  Vol.  38,  no.  3,  pp.  17-19,  32-36. 

1936  Burt,  C.  E.  Amer.  Midi.  Naturalist,  17*,  770-775. 

1936  Parks,  H.  B.,  V.  R.  Cory,  and  others.  Bio/.  Survey  East  Tex.  Big  Thicket 
Area.  Tex.  Acad.  Set.,  pp.  18-20. 

1937  Mulaik,  S.  Copeia,  no.  i,  pp.  72-73. 

1937  Smith,  H.  M.  Herpetologica,  i4,  104-108. 

1938  Burt,  C.  E.  Papers  Mich.  Acad.  Set.  Arts  Letters  ( 1937),  23,  607-610. 
1938    Mulaik,  S.,  and  D.  Sollberger.  Copeia,  no.  2,  p.  90. 

1938  Wright,  A.  H.,  and  A.  A.  Wright.  Trans.  Tex.  Acad.  Set.,  21,  5-35. 

1939  Murray,  L.  T.  Cont.  Baylor  Univ.  Mus.,  no.  24,  pp.  4-16. 

T939  Taylor,  E.  H.  Kans.  Univ.  Sci.  Bull.,  26",  515-518,  529-531,  550-551. 

1940  Taylor,  E.  H.,  and  I.  W.  Knobloch  (see  Arizona). 

1941  Gunter,  G.  Copeia,  no.  4,  p.  266. 

1942  Baker,  R.  H.  Texas  Game  Fish  and  Oyster  Com.,  Bull.  no.  23,  pp.  1-7. 
1942  Gloyd,  H.  K.,  and  H.  M.  Smith.  Bull.  Chicago  Acad.  Set.,  61:J,  231,  232. 
1944  Marr,  J.  C.  (see  General  Wor\s). 

1944    Schmidt,  K.  P.,  and  T.  F.  Smith.  Field  Mus.  Nat.  Hist.,  Zool.  ser.,  29% 
75-96. 

1944  Schmidt,  K.  P.,  and  D.  W.  Owens.  Same,  29",  97-115. 

1945  Smith,  H.  M.,  and  L.  E.  Laufe.  Trans.  Kans.  Acad.  Sci.,  48''*,  325-329. 

1946  Glass,  B.  P.  Copeia,  no.  2,  p.  103. 

1946  Glass,  B.  P.  Herpetologica,  3,  pt.  3,  101-103. 

1946  Netting,  M.  G.,  and  C.  J.  Goin.  Copeia,  no.  4,  p.  253. 

1946  Smith,  H.  M.,  and  B.  C.  Brown.  Herpetologica,  3,  pt.  3,  73. 

1947  Blair,  A.  P.  Copeia,  1947,  p.  137. 

1947  Smith,  H.  M.,  and  B.  C.  Brown.  Proc.  Biol.  Soc.  Wash.,  60,  47-50. 

1947  Smith,  H.  M.,  and  H.  K.  Buechner.  Bull.  Chicago  Acad.  Sci.,  61,  1-16. 

1948  Livezey,  R.  L.,  and  H.  M.  Johnson.  Herpetologica,  4%  164. 
Utah 

1875    Yarrow,  H.  C.  (see  Arizona). 

18843    Cope,  E.  D.  Proc.  Acad.  Nat.  Sci.  Phila.  (1883),  35,  15,  16. 

18840    Cope,  E.  D.  (see  California). 

I9i2a    Van  Denburgh,  J.  (see  Idaho). 

1915    Van  Denburgh,  J.,  and  J.  R.  Slevin.  Proc.  Calif.  Acad.  Sci.,  4th  ser.,  5*, 

99-102. 

1918    Engelhardt,  G.  P.  Copeia,  no.  60,  pp.  77-79. 
1920    Pack,  Herbert  J.  Copeia,  no.  77,  p.  7. 
1922    Pack,  Herbert  J.  Copeia,  no.  102,  p.  8. 
i922b    Pack,  Herbert  J.  Copeia,  no.  107,  pp.  46-47. 
i927a    Tanner,  V.  M.  Copeia,  no.  163,  pp.  54-55. 
i927b    Tanner,  V.  M.  Utah  Acad.  Sci.,  2ist  meeting,  pp.  6-7. 
1928    Tanner,  V.  M.  Copeia,  no.  166,  pp.  23-25. 
1928    Slevin,  J.  R.  (see  British  Columbia). 

Burt,  C.  E.,  and  M.  D.  Burt  (see  Arkansas). 


618  HANDBOOK  OF  FROGS  AND  TOADS 


Burt,  C.  E,,  and  M.  D.  Burt  (see  Arizona). 

1929  Tanner,  V.  M.  Copeia,  no.  171,  pp.  46-52. 

1930  Tanner,  V.  M.  Copeia,  no.  2,  pp.  41-42. 

1931  Tanner,  V.  M.  Proc.  Utah  Acad.  Set.  (Provo),  pp.  159-198. 

1932  Ruthven,  A.  G.  Occ.  Papers  Mus.  Zool.  Univ.  Mich.,  no.  243,  pp.  1-2. 

1932  Taylor,  E.  H.,  and  J.  S.  Wright.  Kans.  Univ.  Set.  Bull.,  2O12,  247-249. 
19333    Burt,  C.  E.  (see  General  Worths). 

1935    Wood,  W.  F.  Copeia,  no.  2,  pp.  100-102. 

1935  Wood,  W.  F.  (see  Lower  California). 
19353    Eaton,  T.  H.,  Jr.  Copeia,  no.  3,  p.  150. 
19355    Eaton,  T.  H.,  Jr.  (see  Arizona). 

1936  Burt,  C.  E.  Amer.  Midi.  Naturalist,  17*,  770-775. 

1936  Cowles,  R.  B.,  and  C.  M.  Bogert  (see  Nevada). 
1938    Hardy,  Ross.  Utah  Acad.  Sci.  Arts  Letters,  15,  99,  102. 
1940    Tanner,  W.  W.  Great  Basin  Naturalist,  i3*  4,  138-139. 
1944    Knowlton,  G.  F.  Copeia,  no.  2,  p.  119. 

Vermont 

1842    Thompson,  Z.  History  of  Vermont,  Natural,  Civil  and  Statistical.  Bur- 

lington. Pt.  I,  pp.  112-113,  119-123. 
1904    Henshaw,  S.  (see  Connecticut). 

1926  Babcock,  H.  L.  (see  Connecticut). 

1931     Loveridge,  A.  Bull.  Boston  Soc.  Nat.  Hist.,  no.  61,  p.  15. 
1938    Fowler,  J.  A.,  and  H.  J.  Cole.  Copeia,  no.  2,  p.  93. 

1940  Trapido,  H.  New  England  Naturalist,  no.  7,  pp.  11-14. 

1941  Moore,  J.  A.  (see  General  Worlds). 
Virginia 

1899    Miller,  G.  S.,  Jr.  (see  District  of  Columbia). 
19153    Dunn,  E.  R.  Copeia,  no.  18,  p.  56. 
I9i5b    Dunn,  E.  R.  Copeia,  no.  25,  p.  63. 
1916    Dunn,  E.  R.  Copeia,  no.  28,  pp.  22,  23. 
1918    Dunn,  E.  R.  Copeia,  no.  53,  pp.  16-22. 
1918    Fowler,  H.  W.  Copeia,  no.  55,  p.  44. 
1920    Dunn,  E.  R.  (see  Alabama). 
1925    Fowler,  H.  W.  Copeia,  no.  146,  pp.  65,  66. 
1925     Brady,  M.  K.  Copeia,  no.  137,  p.  no. 

1927  Brady,  M.  Copeia,  no.  162,  pp.  26-28. 

19293    Burt,  C.  E.,  and  M.  D.  Burt  (see  Arkansas). 
1931     Trautmsn,  M.  B.  Copeia,  no.  2,  p.  63. 

1933  Brandt,  B.  B.,  and  C.  F.  Walker  (see  North  Carolina). 

1937  Brady,  M.  K.  (see  Maryland;  District  of  Columbia). 

1938  Richmond,  N.  D.,  and  C.  J.  Coin.  Ann.  Carnegie  Mus.,  27,  301-304. 
1940    Netting,  M.  G.,  and  L.  W.  Wilson.  Same,  28,  1-8. 

1944  Bartsch,  Paul.  Copeia,  no.  3,  p.  187. 

1945  Hoffman,  R.  L.  Herpetologica,  27~8,  199-205. 

1946  Ackroyd,  J.  F.,  and  R.  L.  Hoffman.  Copeia,  no.  4,  pp.  257-258. 

1946  Hoffman,  R.  L.  Herpetologica,  3,  pt.  4,  141-142. 

1947  Richmond,  N.  D.  Ecology,  281,  53-67. 


BIBLIOGRAPHY  619 

Washington 

1854    Baird,  S.  F.,  and  C.  Girard  (see  Oregon). 

1869    Cooper,  J.  G.  (see  Montana). 

1895    Van  Denburgh,  J.  (see  Idaho). 

1899    Meek,  S.  E.,  and  D.  G.  Elliot.  Field  Columbian  Mus.  Pub.,  no.  31,  Zool, 

sen,  i12,  232-234. 

1899    Stejneger,  L.  Proc.  U.S.  Nat.  Mus.,  2i117S,  899-901. 
191 2c    Van  Denburgh,  J.  Proc.  Calif.  Acad.  Sci.,  4th  ser.,  3,  259-264. 
1916    Dice,  L.  R.  Univ.  Calif.  Pub.  Zool.,  i617,  293-348. 
1918    Ruthven,  A.  G.  Copeia,  no.  53,  p.  10. 

1920  Gaige,  H.  T.  Occ.  Papers  Mus.  Zool.  Univ.  Mich.,  no.  84,  pp.  1-9. 

1921  Blanchard,  F.  N.  Copeia,  no.  90,  pp.  5-6. 

1922  Van  Winkle,  K.  Copeta,  no.  102,  pp.  4-6. 
1928    Slevin,  J.  R.  (see  British  Columbia). 

1931     Noble,  G.  K.,  and  P.  G.  Putnam.  Copeia,  no.  3,  pp.  97-101. 
1931     Slater,  J.  R.  Copeia,  no.  2,  pp.  62-63. 

1933  Svihla,  A.  Copeia,  no.  i,  p.  39. 

J933    Svihla,  A.,  and  R.  D.  Svihla.  Copeia,  no.  i,  pp.  37-38. 

1934  Slater,  J.  R.  Copeia,  no.  3,  pp.  140-141. 

1935  Svihla,  A.  Copeia,  no.  3,  pp.  119-122. 

1939  Brown,  W.  C.,  and  J.  R.  Slater.  Occ.  Papers  Dept.  Biol.,  Coll.  Puget 
Sound  (Tacoma),  no.  4,  pp.  6-31. 

19393    Slater,  J.  R.  Same,  no.  2,  pp.  4-5. 
i939b    Slater,  J.  R.  Herpetologica,  i,  145-149. 

1940  Tanner,  W.  W.  (see  Utah). 

1941  Slater,  J.  R.,  and  W.  C.  Brown.  Occ.  Papers  Dcpt.  Biol.,  Coll.  Puget  Sound 
(Tacoma),  no.  13,  pp.  74-77. 

1942  Slater,  J.  R.  A  Checklist  of  the  Amphibians  and  Reptiles  of  the  Pacific 
Northwest.  Tacoma,  Wash.:  Biol.  Dept.,  College  Puget  Sound.  Mimeo- 
graphed. P.  2. 

1943  Slipp,  J.  W.,  and  G.  C.  Carl  (see  Rtitish  Columbia). 
1945     Schonberger,  C.  F.  (see  Otegon). 

West  Virginia 

i92ga    Burt,  C.  E.,  and  M.  D.  Burt  (see  Aif(ansas). 

1931  Bond,  H.  D.  Copeia,  no.  2,  p.  54. 

1932  Walker,  C.  F.  (see  Ohio). 

1935  Strader,  L.  D.  Proc.  W.  Va.  Acad.  Sci.,  9,  32-35. 

1936  Green,  N.  B.  Same,  10,  80-83. 
1936    Netting,  M.  G.  Same,  pp.  88-89. 

19373    Green,  N.  B.  Herpetologtca,  r,  113-116. 
i937b    Green,  N.  B.  Mag.  Hist,  and  Biog.,  pp.  1-8. 

1938  Green,  N.  B.  Copeia,  no.  2,  pp.  79-82. 

1939  Richmond,  N.  D.,  and  G.  S.  Boggess.  Proc.  W.  Va.  Acad.  Sci.,  12,  57-60. 

1940  Green,  N.  B.  Same,  14,  13. 
1940    Green,  N.  B.  Same,  p.  145. 

1940    Green,  N.  B.,  and  N.  D.  Richmond.  Copeia,  no.  2,  p.  127. 
1940    Llewellyn,  L.  M.  Proc.  W.  Va.  Acad.  Sci.,  14,  148-150. 


620  HANDBOOK  OF  FROGS  AND  TOADS 

1940    Netting,  M.  G.,  and  L.  W.  Wilson.  Ann.  Carnegie  Mus.»2&,  1-8. 

1945    Wilson,  L.  W.  Proc.  W.  Va.  Acad.  Sci.  (1944),  W.  Va.  Umv.  Bull,  no. 

10-1,  pp.  39-41. 
Wisconsin 

1853    Lapham,  I.  A.  Trans.  Wis.  State  Ag.  Soc.  (1852),  2, 366. 

1883    Hoy,  P.  R.  Geol.  Survey  Wis.t  1872-1879,  i,  pt.  II,  425. 

1889      Higley,  W.  K.  Trans.  Wis.  Acad.  Set.  Art  Letters,  1883-1887,  7,  167- 

169,  175. 

1914  Jackson,  H.  H.  T.  Bull.  Wis.  Nat.  His.  Soc.,  new  ser.,  I21'2,  17-20,  22, 
28-29,  35-36. 

1915  Nelson,  T.  C.  Copeia,  no.  19,  pp.  13-14. 

1926    Schmidt,  F.  J.  W.  Copeia,  no.  154,  pp.  131-132. 

1928  Pope,  T.  E.  B.,  and  W.  E.  Dickinson.  Bull.  Pub.  Mus.  Milwaukee,  81, 
1-138. 

1929  Cahn,  A.  R.  Copeia,  no.  170,  pp.  4-6, 

1929  Pope,  T.  E.  B.  Yearboo^  Pub.  Mus.  Milwaukee  (1928),  8,  pt.  I,  177-179, 
181. 

1930  Pope,  T.  E.  B.  Trans.  Wis.  Acad.  Sci.  Arts  Letters,  25,  273,  275,  276,  278, 
279. 

1935  Schmidt,  K.  P.,  and  W.  L.  Necker  (see  Illinois). 

1938  Schmidt,  K.  P.  (see  Illinois). 

1940  Hawkins,  A.  S.  Trans.  Wts.  Acad.  Sci.  Arts  Letters,  32,  63. 

1941  Moore,  }.  A.  (see  General  Worfo). 

1944  Edgren,  R.  A.  J.  Amer.  Midi.  Naturalist,  no.  2,  pp.  495-498. 
Wyoming 

1872  Cope,  E.  D.  (see  Montana). 

1893  Test,  F.  C.  (see  Montana). 

1917  Gary,  M.  North  American  Fauna,  no.  42,  pp.  19,  27,  33. 

1932  Brues,  C.  T.  (see  Nevada). 

1938  Necker,  W.  L.  (see  Illinois). 

1939  Necker,  W.  L.  (see  Illinois). 

MEXICO 

See  Arizona,  New  Mexico,  Texas,  and  General 

Lower  California 

1861     Cope,  E.  D.  Proc.  Acad.  Nat.  Sci.  Phila.,  p.  305. 

1866    Cope,  E.  D.  Same,  18,  312-314. 

1877    Streets,  T.  H.  Bull.  U.S.  Nat.  Mus.,  no.  7,  p.  35. 

1887    fielding,  L.  The  West  American  Scientist,  3-*,  99. 

1895    Van  Denburgh,  J.  Proc.  Calif.  Acad.  Sci.,  2d.  ser.,  5,  pt.  I,  556-560. 

i896a    Van  Denburgh,  J.  Same,  pt.  II,  1004,  1008. 

1899    Mocquard,  F.  Nouv.  arch.  Mus.  dhist.  natur.  (Paris),  4th  ser.,  i,  297- 

299,  334-344- 
1905    Van  Denburgh,  }.  Proc.  Calif.  Acad.  Sci.,  3d  ser.,  Zool.,  4%  1-5,  23. 

1914    Van  Denburgh,  J.,  and  J.  R.  Slevin.  Proc.  Calif.  Acad.  Sci.,  4th  ser.,  4, 
132, 144. 


BIBLIOGRAPHY  621 

19210    Van  Denburgh,  J.,  and  J.  R.  Slevin.  Proc.  Calif.  Acad.  Sci.t  4th  ser.,  n4, 

49-5<>,  53-54- 

1922    Nelson,  E.  W.  Mem.  Nat.  Acad.  Sci.,  i6l,  113. 
1922    Schmidt,  K.  P.  Bull.  Amer.  Mus.  Nat.  Hist.,  46,  art.  XI,  607-634. 

1931  Klauber,  L.  M.  Copeia,  no.  3,  p.  141. 

1932  Linsdale,  J.  M.  Univ.  Calif.  Pub.  2ool.,  38°,  345-354. 

1935  Wood,  W.  F.  (see  Utah). 

1936  Mosauer,  W.  Occ.  Papers  Mus.  Zool.  Univ.  Mich.,  no.  329,  pp.  1-4, 19-21. 
1944    Tevis,  L.,  Jr.  Copeia,  no.  i,  pp.  6,  7. 

1948    Smith,  H.  M.,  and  E.  H.  Taylor  (see  General  Check,  Lists). 


Index 


[Numbers  in  boldface  refer  to  the  pages  of  the  species  account.  1 


Abbott,  C.  C.,  143,  284,  607 
Abbott,  R.  L.,  60 1 
Ackroyd,  J.  R,  618 

Acris,  5,  20,  25,  26,  29,  3$,  50,  67,  77,  87,  88, 
158,  217-28,  229,  232,  242,  246,  306 

—  gryllus  blunchardi,  228 

—  g.  ircpttans,  4,  10,  14,  17.  21,  37,  89,  217, 
218,  220-28,  255,  300,  570 

—  g.  gryllas,  10,  14,  15,  16,  2i,  29,  32,  34, 
37,  50,  78,  87,  89, 198, 217-20,  229,  260,  286, 
468 

Adams,  A.  L.,  591,  603 

Adhesive  disks,  88,  sec  also  Ventral  sucker 

Agassi/,  L.,  253,  548,  591 

Ahl,  K.,  587 

Alabama,  45,  232,  268,  305,  311,  315,  319, 

320,  324,  481,  5 $7,  5*9,  54°>  592-93 
Alaska,  61,  146,  149*  439,  5*>,  544.  549,  593 
Alberta,  164,  180,  250,  252,  481,  544,  59° 
Aldrich,  C.,  60 1 
Alexander,  W.  P.,  609 
Allard,  H.  A.,  212,  215,  216,  598 
Allen,  A.  A.,  255,  609 
Allen,  D.  (L),  605 
Allen,  E.  R.,  340,  472,  597*  598 
Allen,  G.  M.,  347,  480,  544,  607 
Allen,  J.  A.,  315,  603,  606,  610,  615 
Allen,  M.  J.,  238,  269,  472,  496,  538,  539, 

576,  593,  605 
Alligator  frog,  465 
Alhn,  A.  E.,  592 
Amargosa  toad,  84,  154 
Ambystoma,  410;  jeffetsomanum,  26;  macto- 

dactylttmt  520;  maculatunt,  26;  ttgnnum,  26; 

tigrinum  calijorniense,  336 
American  bell  toad,  51,  80,  105 
American  bullfrog,  444 
American  ribbed  toad,  105 
American  toad,  28,  50,  86,  140,  142,  206 
Amphibians,  2;  characters,  2,  3;  orders,  2,  3 
Anderson,  P.,  118,  606 
Anderson  frog,  281 
Anderson's  hyla,  281 

Anderson's  tree  frog,  48,  50,  90,  281,  361 
Anderson's  tree  toad,  281 
Andrews,  E.  A.,  303,  308,  610 
Animal  pole,  36 


Apoda,  2 

Apron,  88;  see  also  Lappet 

Arizona,  7,  10,  32,  58,  61,  73,  101,  111,  112, 
113,  116,  120,  123,  135,  137,  138,  166,  167, 
184,  187,  194,  195*  196,  209,  210,  228,  25?, 
258,  287,  290,  291,  292,  329,  360-68  passim, 
376,  377,  4«"i  444,  447,  448,  473*  475, 
476,  514-19  passim.  553,  554*  555,  576,  577, 
593-94 

Arizona  tree  frog,  287 

Arkansas,  56,  62,  124,  130,  210,  253,  26$, 
272,  311,  344,  347,  353,  450,  497,  540,  594 

Arnold,  L.  W.,  113,  594 

Aronson,  L.  R.,  286,  446,  588,  607,  608,  609 

Airo>o  toad,  156 

Ascaphidae,  3,  20,  32,  55,  80,  81,  104-109 

Astaphus  truci,  4,  12,  16,  18,  20,  31,  33,  34, 
36,  47,  51,  55,  80,  104-10,  413 

Athabaska,  250,  251,  550,  590 

Atkinson,  C.,  600 

Atsatt,  S.  R.,  291,  294 

Authorities'  corner,  79;  see  also  each  species 
account 

Babbitt,  L.  R.,  596 

Babcock,  H.  L.,  588,  596,  603,  604,  607,  614, 
618 

Bahama  Islands,  538,  377,  381 

Bahaman  tree  frog,  377 

Bailey,  J.  R.,  607,  615 

Bailey,  M.  K.,  601 

Bailey,  (O.)  V.,  195,  525,  602,  608 

Bailey,  R.  M.,  405,  408,  409,  60 1,  608 

Baird,  S.  F.,  4,  93,  I02»  i"i  !I3,  M^i  15°, 
192,  220,  240,  242,  244,  247,  263,  269, 
281,  287,  290,  298,  300,  329,  331,  332,  389, 
401,  410,  413,  416,  417,  419,  421,  506, 
509,  511,  516,  520,  524,  533,  535,  536, 
544,  548,  585,  593,  615,  619 

Baird's  toad,  146,  150 

Baker,  R.  H.,  617 

Baldauf,  R.  J.,  614 

Ball,  S.  C.,  596 

Ballou,  W.  H.,  610 

Banded  gecko,  398 

Banta,  A.  M.,  454,  544,  586,  600 

Barbour,  R.  W.,  231,  234,  497,  574,  602 


624 


INDEX 


Barbour,  T.,  2,  4,  5,  104,  116,  172,  184,  244, 
285,  338,  339,  34i,  377.  392,  397,  399»  4<«, 
439,  487,  5<>9,  559,  587.  589*  597,  607 

Barker,  15,  17,  29,  48,  50,  89,  324-2$ 

Barking  frog,  324,  368 

Barking  lizard,  372 

Barnhart,  J.  D.,  413 

Bartram,  W.,  199,  220,  264,  268,  308,  344, 

465.  495.  584 
Bartramian  peeper,  315 
Bartsch,  P.,  309,  618 
Bateman,  G.  C.,  586 
Bawling  frog,  450 
Bel  ding,  L.,  620 
Beldmg's  toad,  192 
Bell  frog,  i,  303,  324 
Bell  toad,  3,  33 
Belly  bumper,  450 
Bcrlandicr,  J.  L ,  511 
Berlandicr's  frog,  481 
Bermuda,  189,  190 
Berndgc,  W.  S.,  587 
Beyer,  G    E.,  410,  602 
Bibbcc,  P.  C.,  252 

Bibron,  G.,  217,  298,  321,  }66,  569,  582,  585 
Bicknell,  E.  P.,  608 
Bird-voiced  hyla,  294 
Bird-voiced  tree  frog,  294 
Bishop,  S.  C.,  212,  213,  535,  536,  537,  602, 

609,  610 
Black,  J.  D.,  594 
Black  chorus  frog,  236 
Black  toad,  84,  176 
Blair,  A.  P.,   216,   589,   598,  601,  602,  604, 

608,  610,  612,  613,  614,  617 
Blair,  W.  F.,  583,  611 
Blanchard,  F.  N.,  497,  520,  599,  600,  601,  602, 

605,  615,  619 
Blatchley,  W.  S  ,  222,  600 
Bloody  nouns,  416,  444 
Bogcrt,  C.  M.,  138,  188,  192,  287,  424,  447» 

476,  506,  575,  589,  593,  595,  607,  618 
Boggess,  G.  S  ,  619 
Boll,  J.,  262,  276,  350,  35? 
Bomare,  J.  C.  U.  de,  584 
Bond,  H.  D.,  619 
Bonnatcrre,  J.  P.,  199 
Bonnet's  frog,  459 
Borland,  J.  N.,  594 
Bosc,  L.  A.  G.,  584 
Boulcnger,  E.  G.,  586 
Boulenger,  G.  A.,  3,  13,  104,  250,  251,  269, 

338,  339,  341,  385,  399*  5°o,  516,  552,  553, 

554»  569,  586 

Boyer,  D.  A.,  222,  227,  606 
Boyle's  frog,  419 
Bradley,  I.  C.,  422,  427,  431,  462,  525 


Brady,  M.  K.,  93,  214,  260,  261,  311,  338, 

339.  495.  597,  603,  618 
Bragg,  A.  N.,  46,  119,  132,  133,  134,  166,  167, 

187,  243,  244,  272,  276,  277,   400,  402, 

496,  502,  509,  510,   578,  583,  587,   588, 

589,  603,  611,  612,  613 
Brandt,  B.  B.,  92,  220,  225,  234,  235,  236, 

268,  324,  572,  598,  610,  618 
Brazil,  1 88 
Breckenndge,  W.  J,  255,  488,  491,  492,  550, 

605 

Breder,  C.  M.,  Jr.,  607,  610 
Breder,  R.  B.,  607,  610 
Breeding,  12-78;  see  also  each  species  account 
Brdim,  A.  E.,  586 
Brcnnan,  L.  A ,  60 1 
Brcukclman,  W.  J.,  602 
Brevicipitidae,  24,  54,  81,  103,  $61-83 
Brimley,  C   S,  217,  218,  219,  224,  228,  235, 

236,  240,  245,  246,  247,  262,  286,  319,  320, 

472,  495,  559,  560,  592,  597,  598,  605,  610 
Brim  ley's  chorus  frog,  92,  234 
Bntcher,  H.  W.,  608 
British  Columbia,  61,  73,  105,  107,  120,  123, 

149,   250,   329,   334,   410,   413,   439,   444, 

447,  481,  520-25  passim,  544,  545.  54^  55i, 

590 

Brocchi,  P.,  367,  585,  586 
Brooking,  W.  J.,  613 
Brown,  A   E ,  586 
Brown,  B   C.,  $53,  480,  617 
Biown,  J.  R.,  591 
Brown,  W  C.,  416,  619 
Brown  garter  snake,  306 
Brownell,  L    W.,  588,  589 
Brues,  C.  T.,  595,  607,  613,  620 
Bryant,  H.  C,  160,  527 
Buechncr,  H.  K.,  617 
Buffon,  G.  L  L  ,  584 
Bttfo.  28,  3r,  33,  50,  77,  83-88,  135-216,  241, 

47T»  5M 

—  alvarius,  TO,  16,  21,  25,  34,  40,  54,  79,  83, 
'35-39?  1 66,  189 

—  amencanus  amencanus,  16,  21,  25,  28,  32, 
34,  40,  50,  59,  78,  86,  139-43,  144,  I52» 
154,  1 60,  1 66,  201,  204,  208,  209,  210,,  216, 

571 

—  amencanus  topet,  21,  86,  140,  143-47 

—  aqua,  190,  191 

—  boreas  Boreas,  13,  16,  18,  21,  25,  31,  41, 
54,  60,  78,  84,  144,  146-51,  155,  159,  160, 
163,  177 

—  boreas  halophtlus,  16,  18,  21,  25,  31,  33, 
34,  41,  60,  61,  78,  84, 149, 150-54, 159, 160, 
163 

—  boreas  nelsonl,  4,  21,  25,  60,  78,  84,  151, 
153-56,  177,  336 


INDEX 


625 


—  cahjotnicus,  4,  16,  21,  31,  34,  40,  54,  79, 
85,  136,  156-61,  171 

—  tanorus,  16,  21,  33,  54,  60,  77,  84,  151, 
160-64,  177,  178.  i79»  43* 

—  cognatus,  16,  17,  18,  21,  25,  34,  40,  58,  78, 
85,  144,  152,  155-60,  163,  164-68,  171,  198 

—  compactilts,  4,  15,  16,  17,  18,  21,  25,  28, 
34,  40,  51,  57,  58,  78,  85,  136,  137, 157,  166, 
167-71,  187,  198,  334,  454,  582 

—  compacttlis  tahformcits,  171 

—  compacttlis  jowlen,  171 

—  tcrmpactihs  speciosus,  171 

—  compactths  woodhousn  t  171 

—  debihs,  4,  16,  18,  21,  25,  31,  54,  77,  87, 
169,  170,  172-75,  184,  185-88 

—  exsul,  4,  1 6,  21,  31,  60,  78,  84,  151,  175-80, 
184,  447 

—  jowlen ,  see  Bufo  w.  jowlen 

—  gtgas,   1 88 

—  hemiophrys,  4,  16,  21,  25,  31,  54,  78,  85, 
144,  180-84,  552 

—  hortibtlts,  192 

—  humeralis,  191 

—  icterus,  191 

—  insidtor,  4,  21,  79,  87,  172,  173, 183-88,  583 

—  fclloggt,  1 86,  1 88 

—  luzarns,  191 

—  lenttgtnostts  amencanus,  see  Bw/o  amen- 
canus 

—  mactilwentns,  191 

—  mcuinus,  16,  21,  86,  144,  188-93 

—  mazatlancnsts,  577 

—  punctatus,  1 6,  21,  25,  28,  34,  37,  41,  51, 
57,  77,  87,  136,  192-97,287 

—  querctcus,  10,  14,  15,  16,  18,  21,  25,  28,  32, 
34,  39,  77,  85,  169,  174,  197-202,  207,  238 

—  speciosus ',  173 

—  terresms,  2,  15,  16,  17,  21,  25,  28,  32,  34, 
41,  50,  58,  59<  77,  85,  137,  M°,  199-202, 
210,  344 

—  valliceps,  16,  21,  25,  28,  34,  41,  51,  58, 

77,  87,  169,  174,  189,  195,  203-206,  242, 
301,  35i,  402,  567 

—  woodhousn  jowlen,  21,  25,  26,  34,  40,  50, 
54»  59,  60,  77,  86,  154,  157,  159,  166,  167, 
206-10 

—  woodhousn  woodhousn,  16,  21,  25,  40,  60, 

78,  86,  152,  154,  157,  159.  166,  167,  206-10 
Bufomdae,  3,  18,  57-61,  81,  83-87,  88,  135-216 
Bullfrog,   12,   13,   19,  30,   32,  96,  305,  416, 

444-50 
Bully,  450 

Bumpus,  H.  C,  480,  614 
Burger,  J.  W.,  614 
Burnett,  W.  L.,  596 
Burns*  meadow  frog,  483 
Burt,  C.  E.,  129,   150,  244,  402,  423,  587, 


588,  593,  594,  595,  596,  599,  601,  602,  605, 
606,  607,  608,  611,  615,  616,  617,  618 

Burt,  M.  D.,  402,  423,  593,  595,  596,  599,  60 1, 
602,  605,  606,  615,  616,  617,  618 

Bushnell,  E.  P.,  588 

Bushnell,  R.  J.,  588 

Butler,  A.  W.,  600 

Cadbury,  J.,  309 

Caecihans,  2 

Cade,  F.  R,  599,  615 

Cahn,  A.  R.,  258,  3241  3^7,  593,  599,  615,  620 

Calhoun,  A  ,  458 

California,  i,  44,  55,  57,  60-62,  74,  101,  105, 
107,  m,  115,  116,  137,  M6,  15<>»  I5-Z* 
154.  156-165  passim,  176,  1 80,  192,  196, 
262,  288-92,  294,  329,  330-36,  410,  41$, 
416-432  passim,  444,  447,  448,  481,  506, 
520,  527,  594-96 

California,  Lower,  see  Lower  California 

California  red-legged  frog,  i,  101,  415-18 

California  toad,  84,  150 

California  yellow-legged  frog,  30,  99,  419-24 

Cambridge  frog,  544 

Camp,  C  L  ,  3,  84,  98,  99,  148,  156,  158,  160, 
163,  171,  287,  329,  419,  421,  424,  428,  429, 
431,  434,  527,  595,  613 

Camp,  R.  D.,  299,  389,  392,  393 

Campbell,  B,  187,  424,  425,  553,  554,  555, 
575,  576,  593 

Campbell,  G.  R.,  597 

Camp's  hog,  10,  95,  373,  389,  398,  416 

Canada,  i,  2,  66,  180,  222,  250,  344,  450,  503, 

504,  533*  549,  590-92 

Canadian  toad,  85,  180-84 

Canyon  toad,  192 

Canyon  tree  trog,  51,  90,  287-94 

Canyon  tree  toad,  i,  32,  287 

Cape  St.  Lucas  spadefoot,  in 

Carl,  G.  C  ,  105,  122,  149,  250,  253,  334,  412, 
413,  447,  520,  522,  590,  606,  619 

Carolina  hyla,  303 

Carolina  narrow -mouthed  toad,  571 

Carolina  toad,  199,  571 

Carolina  tree  frog,  303 

Carolina  tree  toad,  303 

Carpenter  frog,  556 

Carr,  A  F  ,  Jr  ,  46,  129,  ^9,217,219,220,231, 
238,  258,  260-62,  271,  272,  294,  296,  3I7-I9, 
324,  328,  338,  340,  341,  344,  357,  381,  435, 
437,  439,  450,  459,  460,  463,  467,  470,  471, 
49^,  495,  572,  573*  597,  602 

Carr,  K.,  231,  497,  602 

Carter,  H.  A.,  598 

Gary,  M.,  596,  620 

Cascade  frog,  439 

Ca&tenet  frog,  311 


626 


INDEX 


Categories,  3 

Catesby,  M.,  199,  448 

Caudata,  3 

Caulwell,  W.  H.,  211,  286 

Central  America,  298,  302,  516 

Central  Plains  spaclcfoot,  116 

Central  Plains  spadefoot  toad,  116 

Chamberlain,  E.  B.,  219,  224,  234,  235,  236, 
240,  270,  281,  4^,  465,  495,  496,  556,  560, 
614 

Chamberlain,  G.  D ,  547 

Chameleon  hyla,  344 

Chameleon  tree  frog,  i,  344,  347 

Chandler,  A  CM  107,  400-402,  570 

Changeable  tree  toad,  344 

Chapel,  W.  L.,  361,  363,  364,  365,  368,  594 

Chapin,  J,  212,  281,  286,  607,  609 

Charming  toad,  2,  199 

Chase,  H.  B ,  296,  298,  352,  405 

Check  lists,  589 

Chen,  K.  K  ,  598,  602 

Chenu,  J.  C ,  585 

Chorophiltts,  198,  254,  276  (see  also  Psettda- 
ens);  copit,  269;  jeiianim,  260,  jenanim 
brachyphonus,  232;  ntgntus,  260;  occtden- 
talts,  262,  263,  269,  276;  ocnlans,  276; 
ornatus,  262,  269;  septentnonalis,  251,  tn- 
senatus,  230,  240,  251,  256,  260;  tnscrtattts 
tlark.it,  241;  vernicostts,  260 

Chorus  frogs,  i,  14,  229,  244,  361 

Cinereous  frog,  303 

Cinereous  hyla,  i 

Clark,  H.  L.,  604 

Clark,  H.  W.,  228,  258,  483,  591,  600 

Clark,  J.  H.,  190 

Clarke's  chorus  frog,  29,  51,  93,  240 

Clarke's  striped  tree  frog,  240 

Classification,  2-5 

Clausen,  R.  T,  145,  503,  504,  549,  551,  609 

Clay,  W    M.,  6n 

Cliff  frog,  i 

Cnemidophonts,  376,  570 

Coastal  cricket  frog,  217 

Coastal  peeper,  315 

Coat  bet,  14,  324 

Cochran,  D.  M.,  104,  367,  588,  593 

Cockerel!,  T.  D.  A.,  596,  608 

Cockrcll,  V.  L.,  370,  374 

Cold  swamp  frog,  476 

Cole,  H.  J,  618 

Color,  i,  12,  23,  sec  also  each  sptcies  account 

Color  of  throat,  17,  18,  20 

Colorado,  61,  113,  116,  118,  119,  120,  146, 
164,  184,  206,  253,  256,  287,  447,  5°8>  596 

Colorado  River  toad,  10,  83,  135 

Colorado  toad,  135 

Colton,  H.  S.,  258 


Columbian  toad,  146 

Common  bullfrog,  444 

Common  chorus  frog,  244 

Common  frog,  481 

Common  names,  1-2  see  also  each  species  ac- 
count 

Common  spring  frog,  450 

Common  toad,  150 

Common  tree  frog,  344 

Common  tree  toad,  29,  48,  50,  91,  344-47 

Conant,  R.,  311,  504,  556,  560,  597,  599,  608, 
6u,  614 

Connecticut,  220,  228,  500,  504,  512,  596 

Cook,  L ,  158 

Cooper,  I   G.,  138,  329,  505,  595,  606,  619 

Cope,  E.  D,  j,  4,  yo,  91,  93,  116,  120,  137, 
14?,  146,  149,  150,  180,  185,  214,  218,  229, 
250,  232,  240,  241,  244,  245,  246,  258,  260, 
262,  263,  269,  275,  276,  280,  290,  298,  302, 
320,  347,  350-60  passim,  366,  368,  372, 
377,  379,  38' -86  passim,  396,  399,  4Q7»  436, 
473,  474.  476,  479,  492,  497,  502-505  passim, 
508,  511,  512,  51?,  550,  558,  561,  565,  571, 
585,  586,  589-98  pa*sim,  604-20  passim 

Cope's  frog,  556 

Cope's  hyla,  287 

Cope's  spea,  116 

Cope's  tree  frog,  <;r,  347 

Cornell  University,  vni 

Corrmgton,  J.  I),  217,  341,  605,  614 

Cory,  V.  R ,  617 

Cost  A  Rica,  58 

Cott,  H.  B,  588 

Cotton  mouse,  438 

Cotton  rat,  438 

Couch,  D.  N,  174 

Couch's  spadefoot,  i,  27,  83,  111-13 

Coues,  E.,  508,  593,  606,  608,  6 10,  615 

Coventry,  A.  F.,  592 

Cow-bdl  frog,  303 

Cowan,  I  McT  ,  253,  416,  551,  590 

Cowlcs,  R  A.,  476,  593,  607,  618 

Cox,  P.,  591,  592 

Cragm,  F.  W.,  186,  258,  601,  6n 

Crawfish  frog,  402 

Crayfish  frog,   i,  402 

Greaser,  C.  W ,  605 

Cricket  frog,  i,  14,  15,  17,  29,  48,  50,  87,  89, 
217-28 

Cricket  h>la,  217 

Cricket  hylodes,  217 

Cricket  toad,  377 

Croaking,  see  Voice 

Cronise,  T.  F.,  595 

Crotalus,  502 

Cuba,  338,  377,  379,  380,  381,  382 

Culbertson,  116 


INDEX 


627 


Cuvier,  G.  L.  C.  F.  D.,  585 
Cystignathidae,  436 

Cystignathus  fra&lts,  384,  386;  grarilis,  384, 
386;  ornatus,  262 

Danforth,  C.  H.,  605 

Danforth,  S.,  190 

Daniels,  Mrs.  M.  E.,  see  Charlotte  Paine 

Danvers  toad,  210 

Dark  gopher  frog,  537 

Darsie,  Mrs.  R.  F.,  vni 

Daudin,  F.  M.,  191,  266,  268,  269,  270,  275, 

304,  350,  434,  584 
Daughcrty,  Anita,  418,  480 
Davidson,  S.,  366,  367 
Davis,  D.  D.,  599 
Davis,  Mr.  and  Mrs.  L.  L,  394 
Davis,  N.  S.,  Jr.,  102,  400,  402,  407,  589,  599 
Davis,  Norman  W.,  271,  272 
Davis,  W.  T.,  244,  248,  286,  483,  558,  559, 

560,  607,  608,  609 
Deckert,  R.  F.,  129,  202,  214,  215,  216,  220, 

238,  240,  260,  263,  269,  271,  304,  319,  320, 

321,  323»   327,  343,   356,   379,   381,   434, 

437,  464,  586,  597,  608,  614 
Delaware,  220,  248,  309,  311,  500,  504,  59<*-97 
Demerara,  189,  190 
Dent,  J.  N.,  377,  381,  589 
Derrick,  G.  E.,  612 
de  St.  Rcmy,  N.,  478,  592 
Desert  tree  toad,  i,  287 
Desmarest,  M.  E.,  585 
DeSola,  R.,  588 
Development  and  transformation,  75-79,  sec 

also  each  species  account 
Devoe,  A.,  588 
Dicamptodon  ensatus,  107 
Dice,  L.  R.,  578,  593,  601,  619 
Dickerson,  M.  C.,  v,  13, 137,  214,  215,  216,  226, 

256,   286,   342,  356,  401,   402,   418,   464, 

497*   500,   503,   505,   514,   524,  531,   539, 

580,  583,  586 
Dickinson,  W.  E ,  620 
Dieffenderfer,  J.,  180 
Dimorphism,  23 
Discoglossidae,   104 

District  of  Columbia,  7,  246,  308-11,  597 
Ditmars,  R.  L.,  596,  607,  608 
Dixon,  J.  J.,  336,  423,  527,  595 
Dodds,  C.  T.,   191 
Dodge,  N.  M.,  197,  594 
Donaldson,  L.  R.,  105,  606 
Doutt,  J.  K.,  155 
Downs,  A.,  60 1 
Dray  ton,  J.,  417 
Dray  ton's  frog,  416 
Driver,  E.  G.,  126,  588,  604 


Drownc,  F.  P.,  614 

Duellman,  W.  E.,  611 

Duges,  A.,  291,  366,  367,  368,  585,  586 

Dumeril,  A.  M.  C.,  217,  298,  321,  366,  569, 
582, 585 

Duncan,  P.  M.,  584,  586 

Dundee,  H.,  613 

Dunham,  A.  L.,  vni 

Dunkle,  David,  561,  577,  578 

Dunn,  (A.)  M.  T.,  588 

Dunn,  E.  R.,  89,  214,  215,  216,  217,  218,  220, 
223,  224,  232,  308,  379,  381,  588,  592, 
594»  595,  597,  604,  610,  614,  615,  618 

Dury,  R.,  602 

Dusenbcrry,  Mrs.  C.  R.,  367 

Dusky  gopher  frog,  537-40 

Dusky  tree  toad,  90,  353-60 

Dwarf  toad,  197 

Dymond,  J.  R.,  592 


East,  C.  S.,  603 

Eastern  chorus  frog,  17,  29,  93,  244 

Eastern  swamp  cncket  frog,  50,  244 

Eaton,  T.  H.,  Jr.,  290,  291,  476,  514,  588,  594, 

609,  618 

Ecaudata,  2,  3,  104 
Eckert,  J.  E.,  595 
Edgren,  R.  A ,  Jr.,  550,  605,  620 
Edwaids,  W.  E.,  592 
Eggs,  26-46  (see  also  each  species  account); 

laying  process,  31;  mass,  26-31,  36-46,  parts, 

31-36;   seasons,   26    (see  also  Keys,  eggs); 

size,  32 

Ehrhardt,  R.  P ,  105,  108,  109 
Eigcnmann,  C.  H.,  380 
E/aphe,  502;  rosatea,  340,  341 
Elder,  W.  H.,  600 
Eleitthewdiictyhts,  81,  94-95,  366-82 

—  augusti,  4,  95,  3<5<5-68,  374 

—  cactorum,  368 

—  caspan,  382 

—  laticeps,  368 

—  latians,  7,  10,  18,  22,  31,  36,  54,  95,  366, 
367-77 

—  rlcordii  plamrostris,  18,  22,  31,  32,  34,  36, 

54,  95,  37i,  373,  377-8* 
Elliott,  D.  G.,  619 

Ellis,  M.  M.,  150,  253,  256,  447,  596,  604,  608 
Ely,  C.  A.,  192,  589 
Emory,  W.  H.,  615 
Endiley,  J.  R.,  294,  298,  615 
Engelhardt,  G.  P.,  122,  171,  197,  290,  291, 

294,  593,  617 
Engels,  W.  L.,  610 
Engystoma,  568  (see  also  Miciohyla)\  areolata, 

568;  carohncnse,  569,  582;  nucrops,  582; 


628 


INDEX 


Engystoma  (continued) 

olwaccum,  581;  ornatum,  582;  ovalc,  582; 

rugosum,  569,  582;  texcnsc,  569,  581 
Evans,  A.  T.,  604 
Evans,  H.  E.,  610 
Evenden,  F.  G.,  Jr.,  258,  334,  522,  532,  599, 

613 
Evermann,  B.  W.,  228,  258,  483,  524,  525, 

600,  607,  609,  614 
Exner,  I.,  588 
External  gills,  75,  76 


Fannin,  J.,  590 

Fcjervaiy,  G.  J.  de,  104 

Femoral  hyla,  I,  319 

Ferris,  G.  F.,  176,  178 

Fertilization,  75 

Field,  R.  E.,  1 6 

Fisher,  G.  C.,  609 

Fitch,  H.  S.,  416,  419,  422,  423,  595,  613 

Fitzmger,  L.  J.,  229,  383*  584 

Florida,  7,  46,  54,  55,  59,  63,  64,  66,  72,  73, 
74,  124-29  passim,  197*  i99>  2I7»  219,  220, 
236,  238,  240,  248,  258-72  passim,  294, 
295,  296,  303,  304,  306,  315,  317,  318,  319, 
324,  326,  327,  328,  338,  339,  340-44  pass***, 
377,  379.  380,  381,  382,  434-39  passim, 
444,  445,  450,  459-62  passim,  465,  467-72 
passim,  492,  495,  496,  497,  502,  538,  571, 

572,  573»  597-98 
Florida  bullfrog,  459 
Florida  chorus  frog,  93,  258-62 
Florida  frog,  434 
Florida  gopher  frog,  434-39 
Florida  hyla,  324 
Florida  peeper,  91,  3I5'I9 
Florida  tree  frog,  324 
Florida  winter  frog,  258 
Flowers,  Larry,  511,  527 
Fogg,  B.  F.,  603 
Force,  E.  R.,  578,  605,  611 
Fountain,  P.,  220,  465,  598 
Fowler,  H.  W.,  199,  225,  311,  341,  356,  558, 

560,  596,  597,  603,  607,  614,  618 
Fowler,  J.  A.,  324,  603,  618 
Fowler,  S.  P.,  603 
Fowler's  toad,  50,  86,  210-16 
Freeman,  B.,  222,  611 
French  frog,  416 
Freyburger,  W.  A.  Jr.,  614 
Fried  bacon  frog,  303 
Frierson,  L.  S.,  Jr.,  263,  602 
Frogs,  n,  14,  32,  33,  S2,  95-103,  399-5K 

602,  see  also  Ranidae 
Frost,  S.  W.,  609,  614 
Fucrtcs,  L.  A.,  286 


Gadow,  H.,  104,  301,  586,  616 

Gage,  S.  H.,  142,  608 

Gaige,  F.  M.,  394 

Gaige,  H.  T.,  v,  103,  107,  108,  123,  143,  145, 
146,  149,  254,  336,  383,  394,  395,  434, 
5M,  533,  587,  599*  604,  606,  616,  619 

Gaige's  frog,  95,  394 

Gaillard,  Lieut.  D.  D.,  137 

Gannett,  H.,  n 

Garman,  H.,  143,  228,  255,  304,  599,  602 

Carman,  S.,  127,  129,  212,  214,  224,  544,  589, 

615 

Garni,  L.,  276 

Gamier,  J.  H.,  535,  591 

Gasttophtyne,  565,  567,  568,  576  (sec  also 
Micro hyla) ;  areolata,  568;  carolinensis,  569, 
581;  elegans,  577,  581;  ohvacea,  565,  569, 
581;  texcnsis,  576,  577;  ttsta,  576 

Gebhard,  J.,  Jr.,  608 

Geikie,  J.  C.,  347,  450,  586 

General  works,  584-89 

Gentry,  G.,  615 

Geological  speculations,  5-10 

George,  I.  D.,  602,  605 

Georgia,  65,  67,  124,  140,  199,  201,  202,  212, 
219,  220,  223,  227,  234,  235,  236,  238,  240, 
264-69  passim,  294,  303,  304,  306,  311, 
315,  3i8»  320-28  passim,  344,  356,  434-39 
passim,  459,  461-65  passim,  469,  470,  471, 
481,  492,  494,  495,  496,  504,  540,  556,  559, 
560,  580,  598 

Georgia  tree  frog,  324 

Getrhonotus  infer  nalis,  372 

Gcssing,  W.,  Jr.,  602 

Giant  toad,  135,  188 

Giant  tree  frog,  89,  324,  337-41 

Gibbs,  M.,  604 

Gier,  H.,  409,  509,  510,  511,  588,  611 

Gilbert,  E.  S ,  608 

Gilbert,  P.  W.,  609 

Gilmore,  R.  J.,  118,  119,  120,  596 

Giovanolh,  L.,  602 

Girard,  C,  10,  I35,  137,  150,  171,  172,  184, 
185,  186,  192,  206,  262,  263,  269,  287,  329, 
33i,  332,  399,  401,  410,  413,  416,  417,  516, 
520,  524,  569,  582,  585,  615,  619 

Girard's  toad,  135  • 

Glass,  B.  P.,  342,  617 

Gloyd,  H.  K.,  226,  405,  408,  410,  588,  594, 
599,  601,  617 

Gmelm,  C.  C.,  584 

Goergcs,  Mrs.  F.  C.,  vm 

Coin,  C.  J.,  46,  102,  216,  220,  304,  311,  317, 
377,  381,  382,  399,  537,  538,  539,  540,  588, 
589,  593,  597,  598,  602,  605,  617,  618 

Gopher  frog,  i,  15,  30,  49,  75,  102,  402,  434 

Gopher  turtle,  434,  539 


INDEX 


629 


Gordon,  K.,  46,  107,  410,  423,  447,  520,  527, 

532,  533.  613 
Gorsuch,  D.  M.,  138 
Graf,  W.,  613 
Grant,  C.,  60 1 

Grant,  R.,  146,  476,  504,  533,  535,  550 
Grass  frog,  481 
Grasshopper  frog,  i 
Grave,  B.  H.,  600 
Gravenhorst,  J.  L.  C.,  585 
Gray,  I.  E ,  610 
Gray,  J.  E.,  585 
Gray  toad,  199 

Great  Basin  spadcfoot  toad,  120 
Great  Plains  toad,  17,  85,  164 
Green,  H.  T.,  609 
Green,  N.  B.,  124,  231,  234,  619 
Green  and  yellow  tree  toad,  281 
Green  bullfrog,  459 
Green  frog,  19,  49,  53,  98,  45<>-54 
Green  toad,  i,  172 

Green  tree  frog,  29,  30,  48,  89,  303-308 
Greenback,  465 
Griffiths,  E.,  585 
Gnnneil,  J.,  148,  154,  160,  163,  196,  287,  329, 

336,  416,  419,  423,  424,  434,  527,  595,  596 
Gronbcrger,  S.  M ,  597 
Guntcr,  G.,  617 
Gunther,  A.,  188,  302,  383,  585,  586 


I- labor,  J  M.,  213,  246 

Haber,  V.  R.,  213,  246,  308,  587 

Habitat,  10,  see  also  each  species  account 

Hadsell,  L.  T.,  116,  418,  432 

Hahn,  W.  L.,  600 

Haldeman,  S.  S.,  613 

Hall,  E.  R.,  473 

Hall,  H  M.,  595 

Hallman,  T.,  272,  439,  597 

Hallowell,  E.,  304,  332,  575,  578,  582,  585, 

594,  597,  601,  608,  613,  615 
Hallowell's  tree  frog,  303 
Haltom,  W.  L.,  593 
Hamilton,  W.  J.,  Jr.,  viii,  362,  609 
Hammond's  spadefoot,  10,  27,  76,  77,  82,  113, 

116,  124 

Hammond's  spea,  113 
Hankinson,  T.  L ,  228,  599,  604 
Hardy,  G.  A.,  590 
Hardy,  R.,  122,  618 
Hargett,  C.  W.,  604 
Hargreaves,  C.  C.,  598,  602 
Harlan,  R.,  124,  584,  585 
Harper,  R,  91,  93,  228,  229,  238,  240,  260, 

268,  272,  286,  294,  296,  315,  317,  318,  319, 

321,  356,  357,  377,  438,  439,  470,  558,  573» 


587,  588,  589,  591,  592,  597,  598,  602,  605, 

610,  614 

Hartman,  F.  A.,  167,  410,  601 
Hartweg,  N.,  587 
Harvey,  G.  W.,  514,  554,  555 
Harwell,  C.  A.,  160,  429 
Harwood,  P.  D.,  401,  402,  570 
Hassler,  W.  G ,  308,  572,  603 
Hatfield,  D.  M.,  155 
Hatt,  R.  T.,  605 
Hawkins,  A.  L.,  620 

Hay,  O.  P.,  214,  215,  253,  257,  513,  599,  600 
Hay,  W.  P.,  246,  311,  586,  597,  603 
Hay  frog,  253 
Haydcn,  F.  V.,  513,  606,  615 
Hazel  tine,  K.,  595 
Hazzard,  A.  L.,  105 
Hecht,  B.  L.  M.,  568,  574,  575,  589 
Hecht,  M.  K.,  103,  568,  574,  575,  589 
Heckschcr's  frog,  465-72 
Ilcmze,  A.  A.,  222,  227,  606 
Heller,  J.  A.,  609 

Helocaetes  tlarfyi,  240,  242;  jcnarum,  247 
Henderson,  J.,  150,  253,  256,  447,  596 
Hennmg,   W.    L.,    606 

Henshaw,  H.  W,  145-46,  364,  422,  595,  606 
Henshaw,  S.,  596,  603,  604,  607,  614,  618 
Hermit  spadefoot,  124;  toad,  124 
Hcrrara,  A.  L ,  586 
Herring  hoppers,  481 
HcfctanJna  jormosa,  10 
Hibbard,  C.  W.,  602 
Higlcy,  W.  K.,  620 
Hill,  T.,  603 
Hilton,  J.  W  ,  576>  577 
Hinckley,  M.  H,  v,  171,  210,  313,  314,  315, 

347,  542,  543.  544,  604 
Hitchcock,  E.,  603 
Hobbs,  K.  L.,  593 
Hoffman,  R.  L ,  347,  618 
Holbrook,  J.  E.,  in,  140,  146,  197,  199,  202, 

212,  219,  263,  264,  268,  271,  304,  305,  321, 

323,  342,  374,  495,  57i,  574»  58fi  582,  585, 

598 

Holbrook's  chorus  frog,  268 
Holbrook's  spadefoot,  10,  124,  128,  129 
Holt,  E.  G.,  593 
Hoopes,  1 ,  588,  603,  604 
Hoosier  frog,  402,  407,  533 
Hoover,  T.  S.,  335,  418,  423 
Hop  toad,  140,  199 
Hopson,  R.  E.,  107,  335,  414,  532 
Horny  excrescences,  23-25,  80 
Hough,  F.  B.,  608 
House  frog,  i 

Howe,  R.  H.,  Jr.,  544,  552,  586 
Hoy,  P.  R.,  620 


630 


INDEX 


Hubbs,  C.  L.,  176,  180,  216,  458,  599,  600, 

605 

Hudson,  G.  E.,  225,  606 
Hudson  Bay,  7,  59,  6*3,  66,  143,   145,  250, 

476,  533,  544,  548 
Hudson  Bay  American  toad,  143 
Hudson  Bay  toad,  86,  143 
Hughes,  E.,  600 
Humphrey,  R.  R.»  138 
Humphreys,  J.  T.,  610 
Hurter,  J.,  225,  228,  253,  262,  263,  280,  410, 

539,  594,  595,  599,  605 
Hurter 's  solitary  spadefoot,  130 
Hurter's  spadefoot,  83,  130 
Hyla,  13,  14,  20,  25,  26,  29,  31,  50,  77,  87-91, 

94,  159,  232,  281-365,  571 

—  affinis,  290 

—  andersomi,  14,  16,  20,  22,  26,  31,  34,  36, 
48,  50,  72,  78,  90,  281-86,  343,  361,  504 

—  aremcolor,  22,  26,  31,  32,  34,  37,  51,  73, 
74,  78,  90,  n6,  287-94,  329,  428,  514 

—  avivoca,  i,  16,  22,  26,  31,  54,  79,  91,  294-99, 
352,  355,  356,  357,  359,  3^o 

—  baudinu  batidtmt,  16,  22,  26,  31,  54,  78,  90, 
289,  298-302,  340 

—  brunnea,  339 

—  ctnerca  ctnetea,  7, 16,  20,  22,  26,  29,  31,  34, 
38,  48,  50,  72,  78,  89,  283,  286,  296,  300, 
303-308,  309,  310,  311,  326,  342,  343,  463, 
470,  471,  500,  538,  571 

—  cmcrca  evittata,  4,  7,  20,  22,  26,  31,  54,  89, 

303,  305,  307-" 

—  cinerea  semifasciata,  304 

—  cntcifer  baitianiana,  4,  QT,  312,  3I5'I9 

—  cructfer  cntufer,  16,  21,  26,  29,  31,  34,  37, 
48,  50,  68,  69,  78,  91,  231,  232,  234,  246, 
254,  271,  296,  311-16 

—  eximia,  361,  362 

—  jemoralis,  14,  15,  16,  18,  22,  26,  29,  32,  33, 
34,  38,  48,  5°,  73,  78,  90,  198,  229,  258, 
294,  295,  319-249  344,  350,  352,  355 

—  femoralts  chrysoscelts,  350,  353 

—  ilavlgtda,  342 

—  gratiosa,  4,  15,  16,  17,  20,  22,  26,  29,  33, 
34,  37,  48,  50,  7*»  72,  78,  89,  198,  282,  319, 
324-28,  373,  438,  538 

—  lafrcntzi,  361,  362 

—  laterals,  304,  341 

—  nebulosa,  262 

—  ocularis,  269 

—  phaeocrypfa  o  gee  fan  f is,  360 

—  picfertngi,  246,  254,  271,  see  H.  cructfer 

—  rcgilla,  16,  22,  26,  31,  34,  46,  73,  7$,  94, 
156,  159,  164,  262,  267,  287,  288,  291,  294, 

3^9-37*  423,  454,  455 

—  scapttlaris  hypochondriaca,  262 

—  scmijasdata,  304 


—  septcntrionalis,  10,  16,  18,  22,  24,  25,  79, 
89,  282,  337-41 

—  squirclla,  16,  21,  26,  29,  34,  36,  48,  50,  74, 
78,  90,  198,  232,  235,  306,  320,  321,  322, 
323,  341-45,  350 

—  van  vlictii,  298,  300 

—  vcrstcolor  chrysoscelis,  4,  16,  22,  31,  54,  91, 
174,  289,  347-53 

—  verstcolor  phaeocrypta,  4,  22,  54,  90,  353-6o 

—  vets/color  sandersi,  353 

—  versuolor  venicolor,  16,  22,  26,  29,  31,  34, 
38,  48,  50,  74,  78,  91,  232,  242,  246,  278, 
286,  287,  289,  290,  294,  296,  320,  323,  344- 
47>  348,  349,  350,  351,  352,  353,  355-6*0, 
57is  583 

—  virtdis,  304 

—  wrishtorum,  4,  22,  31,  54,  78,  94,  360-65 
Hyhdae,  3,  18,  20,  21,  22,  25,  33,  67-84,  82, 

87-94,  217-365 
Hylodes  ocularis,  266 
Hypopathus,  561;  cuneus,  12,  24,  35,  38,  54, 

78,  103,  204,  561-67 

Idaho,  7,  60,  105,  108,  116,  120,  206,  255,  329, 
330,439,444,447,598-99 

Illmgworth,  J.  T.,  588 

Illinois,  26,  72,  212,  224,  253,  255,  256,  303, 
304,  355,  356,  358,  359,  402,  405,  406,  407, 
409,  410,  481,  496,  497,  505,  510,  511,  513, 
548,  57i,  599-6oo 

Indiana,  54,  64,  74,  222,  253,  257,  341,  356, 
359,  402,  407,  408,  409,  481,  495,  496,  497, 
502,  505,  510,  513,  548,  571,  600-601 

Ingram,  W  M ,  152,  335,  588,  609 

Internal  gills,  76 

Iowa,  60,  206,  2TO,  227,  450,  511,  601 

Jackson,  H.  H.  T.,  620 

Jackson,  H.  W.,  145,  620 

Jamaica,  377,  381 

James,  J.  E.,  600 

James,  J.  F.,  611 

James  Bay,  143,  145,  476,  549 

Janes,  R.  G.,  588,  609 

Jellison,  W.  J.,  105,  329,  330,  522,  606  4 

Jelly  envelopes,  33,  34,  35,  75,  see  also  Keys, 

eggs 

Jcwett,  S.  G.,  Jr.,  336,  410,  520,  613 
Joe  Brown  frog,  459 
Johansen,  F.,  591 
Johnson,  H.  M.,  617 
Johnson,  Harlen,  123 
Johnson,  J.  E.,  617 
Jones,  J.  M.,  591 
Jones,  R.  W.,  612 
Jopson,  H.  G.  M.,  614 


INDEX 


63' 


Jordan,  D.  S.,  406,  585,  587 
Jug-o'-rum,  444 

Kalm,  Peter,  483,  584 

Kansas,  60,  63,  116,  118,  119,  140,  164,  166, 
167,  184,  186,  187,  206,  208,  209,  222,  223, 
226,  240,  241,  243.  253,  257,  311,  344,  355. 
402,  404,  405,  407,  408,  444,  45o;  48i,  5°5, 
509,  510,  511,  578,  581,  583,  601-602 

Kapp,  Mrs.  K.,  viu 

Kauffeld,  C.  R,  137,  138,  187,  496,  497,  500, 

503,  504.  505,  587,  594,  609 
Keewatm,  250,  590 
Keim,  T.  D.,  603,  614 
Keller,  J.  G.,  553,  555,  608 
Kellogg,  R.,  1 1 8,  119,  174,  1 86,  187,  188,  189, 

206,  302,  365,  366,  367,  373,  384,  503,  5M. 

516,  587,  593,  606 
Kelly,  H.  A.,  603 
Kentucky,  7,  229,  231,  233,  234,  253,  294,  402, 

481,  492,  497,  505,  571,  602 
Kerr,  R.,  584 

Key  West  spadetoot,  8?,  127 
Key  West  tree  frog,  10 
Keys:  eggs,  36-46;  families  and  species,  80-103; 

tadpoles,  54-74 

Kilby,  J.  D.,  303,  306,  308,  434,  495,  598 
King,  F.  W.  (see  W.  King),  nj,  116,  138,  167, 

291,  366,  515 
King,  H.  D ,  v 
King,  W.,  610,  615 
Kmgsley,  J.  S.,  586 
Kirkland,  A  H.,  143,  603 
Kirn,  A.  J.,  150,  174*  243,  276,  279,  280,  351 
Kirsch,  P   H.,  600,  611 
Kirtland,  J.  P.,  610 
Klauber,  L.  M.,  136,  156,  157,  158,  159,  165, 

171,  288,  331,  332,  416,  417,  424,  426,  428, 

593.  595,  607,  621 
Klots,  A.  B.,  283,  285,  286,  608 
Klugh,  A.  B.,  591 
Knepp,  T.  H.,  614 
Knight,  H.  H ,  320 
Knobloch,  I.  U.,  594,  617 
Knowlton,  G.  P.,  599,  618 
Knox,  C  B.,  547 

Koster,  W.  J.,  366,  374,  375,  376,  593,  608 
Kuntz,  R.  E.,  187,612 

Labia,  47,  50,  51,  53,  76 

Labial  papillae,  47,  50,  51,  53 

Labial  teeth,  47,  50,  51,  53,  76 

Labrador,  143,  145,  481,  544,  59°-9i 

Lacepcde,  B.  G.  E.  de  V.,  584 

Lachner,  E.  A.,  610 

Lake  frog,  459 

Lampropeltis  triangulum  temporalis,  504 


Land  frog,  199 

Land  toad,  199 

Lapham,  I.  A.,  620 

Lappet,  23,  88 

Lateral  lines,  47 

Latham,  R.,  609 

Latrcille,  P.  A.,  199,  319,  341,  450,  584 

Latreillefs  toad,  199 

Laufe,  L.  E ,  617 

Laurcnti,  J.  N.,  104,  135,  584 

Least  swamp  frog,  264 

Le  Conte,  J.,  vn,  217,  226,  236,  238,  267,  305, 

321,  324,  327,  344.  434,  436,  476,  479,  54°, 

542,  573»  58i,  584,  585,  598 
Le  Contc,  J.  L.,  262 
Le  Contc's  chorus  frog,  236 
Lc  Conte's  frog,  416 
Le  Conte's  leopard  trog,  476 
Leffingwell,  D.  J.,  330 
Leighton,  H.,  481 
Lentz,  M.  J.  RM  594 
Leonard,  A.  B.,  60 1,  611 
Leonard,  M.  J-.,  190 
Leopard  frog,  476,  481 
Lcptodactylidae,  3,  22,  25,  81,  82,  94-96,  97, 

366-98 

Lfptodactyhts  albdabns,  382,  384,  385 

—  fiagtlis,  384 

—  giatths,  384 

—  labmlis,  22,  371,  383-88 
Lett,  W  P ,  591 

Leums  J.,  585 

Life  Zones,  6,  8,  9,  n,  140,  15",  151,  156,  206, 

281,  311,  315,  319,  329,  360,  }6r,  416,  481, 

500,  503,  504-506,  509,  514 
Linnaeus,  C.,  188,  399 
Linsdale,  J.  M.,  101,  121,  122,  154,  155,  156, 

171,  210,  222,  336,  416,  418,  423,  447,  454, 

457-59,  473,  475,  5<>6,  519,  527,  532,  553, 

555,  595,  596,  599,  601,  607,  608,  621 
Lmslcy,  J.  H.,  596 
Liopelmidae,  104 
lithodytes  lattans,  280 
Lttona  occidenlahs,  262 
Little,  h  S.,  Jr  ,  287,  553,  555,  594,  608 
Little  chorus  frog,  10,  15,  29,  48,  50,  92,  264 
Little  green  toad,  87,  172-75 
Littleford,  R.  A,,  216,  603 
Liu,  C.  C.,  16,  587 
Live/cy,  R.  L.,  32,  34,  36,  107,  138,  159,  414, 

422,  423,  432,  531,  589,  617 
Llewellyn,  L.  M.,  619 
Lockmgton,  W.  N.,  595 
Lodmg,  H.  P.,  539,  592 
Locnnbcrg,  E.,  199,  574,  597 
Logier,  E.  B.  S.,  105,  107,  121,  124,  146,  215, 

520,  524,  525,  589,  590,  591,  592 


INDEX 


Long-footed  frog,  416 

Loonus,  R.,  60 1,  606 

Loskiel,  G.  H.,  347,  454.  483.  584 

Louisiana,  10,  58,  59,  63,  64,  65,  66,  72,  197, 
199.  203,  214,  217,  222,  223,  225,  240,  253, 
264,  294,  296,  298,  311,  319*  3^4)  352,  353, 
358,  399,  4°5»  45°,  459,  464»  4^5*  49^,  497, 
502,  537,  538,  602-603 

Lovendgc,  A.,  587,  618 

Lower  California,  44,  57,  73,  in,  113,  I5°» 
156,  160,  192,  194,  196,  287,  329,  416, 
620-21 

Lucania  ommata,  10 

Lynn,  W.  G,  302,  377,  381,  589 

Mabie,  W.  B.,  610 

McAtee,  W.  L.,  597,  600,  603,  607 

McCauley,  R.  H.,  Jr.,  338,  603 

McCiung,  C.  E.,  588 

MacCoy,  C.  U.,  593 

McFarland,  R  W.,  539 

Mackay,  A.  H.,  591 

McKee,  E.  D.,  258,  287,  593 

Mackenzie,  180,  250,  251,  481,  544,  550,  59* 

McLellan,  M.  E.  (Mrs.  Davidson),  367 

McManus,  I.,  614 

Maine,  74,  344,  444,  5*2,  513,  544>  547*  593 

Mandible,  47,  76 

Manitoba,  140,  180,  182,  252,  311,  344,  550, 

591 

Manitoba  toad,  180 

Mansueti,  R.,  603 

Manvillc,  R.  H.,  603 

Marimon,  S ,  595 

Marine  toad,  86, 188 

Mark,  E.  L.,  189,  190 

Marnock,  G.  W.,  276,  298,  350,  370,  373,  374, 

396 

Marnock's  frog,  95,  96,  373,  395-98 
Marr,  J.  C.,  210,  398,  424,  589,  596,  602,  606, 

613,  617 
Marsh  frog,  476 
Marsh  tree  frog,  303 
Marshall,  B.  C.,  396 
Martin,  F.,  616 
Maryland,  72,  220,  229,  232,  248,  308,  309, 

319,  324,  448,  500,  504,  540,  542,  543,  544, 

549,  571,  572,  593 
Massachusetts,  55,  60,  124,  126,  313,  314,  509, 

512,  603 

Matalas,  B.  L,  103,  568,  574,  575,  5^9 
Mattern,  E.  S.,  614 
Mattern,  W.  L,  614 
Maynard,  E.  A.,  609 
Meadow  frog,  3,  49,  102,  481-83 
Mearns,  E.  A.,  137,  138,  462,  514,  593,  595, 

608,  616 


Meek,  S.  E.,  591,  619 

Merrcm,  B.,  584 

Mernam,  C.  H.,  6,  505 

Merryday,  M.  P.,  598 

Metamorphosis,  sec  Transformation 

Metcalf,  Z.  P.,  587 

Mexican  cliff  frog,  95,  366 

Mexican  frog,  98,  553-55 

Mexican  toad,  203 

Mexican  tree  frog,  90,  298 

Mexico,  5,  57,  58,  60,  61,  64,  74,  in,  113, 
164,  167,  171,  172,  174,  184,  186,  187,  188, 
191,  192,  193,  203,  287,  301,  302,  360,  383, 
384,  481,  509,  511,  513,  516,  553,  562,  577, 
578,  620-21 

Michigan,  210,  253,  254,  255,  409,  444,  497, 
512,  544,  548,604-605 

Mtcrohyla,  5,  7, 14,  26,  27,  53, 174,  561, 568-83 

—  areolata,  5,  24,  54,  103,  562,  568-72 

—  caioltneims,  14,  15,  16,  17,  24,  27,  32,  33, 
35,  37.  48,  53,  54,  77,  i<>3,  213,  562,  568, 
569,  57°,  57^-74*  575,  578-82  passim,  583 

—  elegant,  577 

—  mazatlanensis,  5,  103,  562,  574-77 

—  ohvacea,  12,  16,  24,  27,  33,  35,  39,  54,  77, 
103,  174,  185,  242,  306,  351,  509,  562,  569, 
57^,   574,   575,  577,  578-83 

—  rugosum,  569 

—  texensts,  16,  570 

—  tuta,  577 
Miles,  M.,  604 

Miller,  A.  H,  156,  158,  596 

Miller,  D.  C.,  600 

Miller,  G.  S.,  Jr.,  272,  308,  309,  310,  311,  533, 

597,  618 

Miller,  L.,  1*56,  596 
Miller,  R.  R.,  197,  458,  596 
Miller,  W.  D.,  212,  214,  215,  216,  281,  607, 

609 

Miller's  tree  fiog,  89,  307-11 
Mink  frog,  2,  19,  49,  53*  98,  533-4® 
Minnesota,  58,  66,  74,  140,  164,  250,  252,  253, 

3".   344,   353,    355,  35&>   359,   444,   45°, 

483-92  pass*m,  500,  502,  504,  511,  533,  535, 

540,  544,  550,  55i,  552,  605 
Mississippi,  45,  199,  225,  229,  232,  236,  238, 

262,  269,  306,  308,  311,  315,  34i,  4°2,*497» 

512,  538,  539,  540,  605 
Mississippi  River,  303,  402,  492,  496,  497,  504, 

510,  582 
Missouri,  60,  1 1 8, 124,  164,  206,  210,  222,  225, 

227,  303.  402,  481,  497,  539,  578,  605-606 
Mitchell,  J.  D.,  103,  568 
Mitchell's  narrow-mouthed  toad,  103,  568-71 
Mittlcman,  M.  B.,  356,  357,  409,  509,  510,  511, 

588,  589,  602 
Mocquard,  F.,  138,  366,  367,  586,  616,  620 


INDEX 


633 


Moesel,  J.,  609 

Mohr,  C.  £.,  247,  614 

Monks,  S.  P.,  608 

Montana,  7,  56,  61,  105,  118,  119,  146,  164, 

206,  250,  253,  329,  330,  508,  509,  513,  520, 

522,  525,  606 
Moore,  B.,  592 
Moore,  C.  B.,  609 
Mooie,  G.  A.,  46,  120,  167,  210,  228,  243,  280, 

612 

Moore,  H.  F.,  464 
Moore,  J.  A  ,  4^7,  497,  503,  504,  505,  511,  588, 

589,  598,  604,  605,  608,  609,  618,  620 
Morgan,  A.  H.,  588 
Morse,  M.,  253,  611 
Mosauer,  W.,  608,  616,  621 
Mottled  meadow  frog,  488 
Mountain  chorus  frog,  94,  229 
Mountain  toad,  146 
Mouths  of  tadpoles,  47,  50,  51,  53 
Mulaik,  D,  388 

Mulaik,  S.,  366,  385,  387,  388,  561,  617 
Mulford,  F.  L.,  8,  9 
Muller,  J.  W.,  585 
Munz,  P.  A.,  196 
Murray,  L.  T.,  292,  294,  617 
Muscle  segments  (myotomcs),  47,  75,  76 
Musgrove,  M.  E.,  593 
Myers,  G.  S.,  104,  108,  158,  163,  176-80,  214, 

215,  216,  311,  407,  408,  409,  447,  587,  595, 

596,  598,  600,  607-10  passim 

Nairow  mouth  frog,  571 

Narrow  mouth   toad,   i,   15,   17,  27,  48,  53, 

571-74 

Narrow-mouthed  frog,  33,  571 
Narrow-mouthed  toad,  3,  12,  14,  32,  76,  77, 

81,  103,  571 
Nash,  C.  W.,  591 
Nattix  rhomlnjera,  206 
Nebraska,  225,  226,  227,  253,  444,  571,  578, 

606 

Nebulous  toad,  28,  51,  87,  203,  571 
Necker,  W.  L.,  394,  548,  588,  599,  600,  605, 

615,  620 

Necdham,  J.,  420 
Needham,  P.,  292 
Nail,  W.  T.,  360,  598,  614 
Nelson,  E.  W.,  621 
Nelson,  J.,  607 
Nelson,  T.  C.,  620 
Netting,  M.  G.,  102,  214,  215,  216,  224,  232, 

399,  537,  538,  539,  54°,  587,  5^9,  598,  602, 

614,  617-20  passim 
Nevada,  7,  42,  60,  66,  101,  120,  121,  122,  137, 

146,  150,  154,  155,  156,  164,  167,  192,  206, 

287,  329,  336,  444,  447,  448,  454-59,  473, 


474,  475,  47<>»  481,  506,  512,  513,  514,  519, 

527-33  passim,  606-607 
Nevada  frog,  19,  51,  101,  453'59 
Nevada  spotted  frog,  30,  100,  527-33 
New  Brunswick,  140,  311,  450,  476,  533,  549, 

59i 

New  England,  12,  66,  444,  445,  502,  504,  535 
New  Hampshire,  210,  607 
New  Jersey,  14,  42,  67,  72,  211,  212,  220,  225, 

227,  244,  245,  247,  281-86  passim,  450,  481, 

488,  496,  500,  502,  504.  505,  510,  556,  558, 

559,  560,  607-608 
New  Mexico,  10,  61,  113,  116,  137,  184,  187, 

206,  222,  253,  360,  364,  368,  374,  376,  444, 

447,  502,  508,  553,  608 
New  Mexico  spea,  113 
New  York,  66,  127,  142,  210,  212,  220,  228, 

244,  254-58  passim,  314,  315,  347,  445,  44^, 

452,  480-83  passim,  497,  500,  504,  506,  514, 

533-37  passim,  541-44  passim,  608-10 
Newcome,  W.  A.,  590 
Newfoundland,  481,  591 
Nicholas,  G.  A  ,  405,  409 
Nichols,  A.,  1 27,  603 
Nichols,  J  T.,  609 
Niedcn,  F.,  104,  587 
Noble,  G   K.,  2,  104,  190,  268,  285,  286,  308, 

3&3«  572,  586,  587,  588,  603,  607,  619 
Noble,  R.  C.,  285,  286,  607 
North  American  bullfrog,  444 
Noith  Carolina,  14,  59,  63,  72,  197,  199,  213- 

20  passim,  224,  234,  235,  236,  244-48  passim, 

264,   281,   286,  319,   324,   346,  360,  434, 

465,   481,   495,   504,   540,   572,   573,   574, 

610 
North  Dakota,  116,  180,  181,  182,  183,  250, 

2^1,  252,  253,  ^57,  489,  508,  546,  552,  610 
Northern  chorophilus,  250 
Northern  crayfish  frog,  400 
Northern  frog,  535 
Northern  gopher  frog,  15,  102,  402-10 
Northern  leopard  frog,  495 
Northern  pseudacris,  250 
Northern  spring  peeper,  250 
Northern  stnpul  tree  frog,  92,  250 
Northern  toad,  140,  146 
Northern  tree  toad,  344 
Noithern  wood  frog,  12,  99,  544-52 
Noith  western  toad,  84,  146 
Norton,  A.  H.,  535 
Nova  Scotia,  62,  140,  535,  540,  549,  591 


Oak  frog,  197 
Oak  toad,  10,  15,  28,  48,  ! 
Obrccht,  C.  B.,  614 
O'Donoghue,  C.  H.,  591 
Odor,  2 


197,  434 


634 


INDEX 


Ohio,  7,  45,  63,  124,  210,  212,  229,  231,  253, 
255,  402,  409,  610-11 

Ohio  chorus  frog,  229 

Oklahoma,  46,  56,  63,  64,  116,  119,  132,  140, 
164,  166,  167,  187,  206,  222,  240,  241,  243, 
244,  253,  272,  276,  277,  344,  399,  4oo,  402, 
444,  450,  496,  497,  502,  505,  510,  511,  578, 
583,  611-13 

Olive-colored  frog,  481 

Oliver,  J.  A.,  138,  188,  192,  506,  575,  576, 
587,  589,  607 

Ontario,  7,  62,  143,  145,  253»  444,  44$,  447, 
476,  478,  481,  533,  536,  540,  544,  549, 
591-92 

Oppel,  M.,  584 

Oregon,  7,  44,  46,  61,  62,  105,  107,  149,  206, 
329,  33i,  332,  334,  335,  336,  410,  4«i  4*3, 
414,  419,  422,  423,  439,  441,  444,  447, 
448,  481,  513,  520,  522,  525,  527,  528,  529, 

550,  532,  535,  6*3 
Oregon  red-legged  frog,  101,  410-16 
Ornate  chorus  frog,  94,  267-72 
Ornate  swamp  frog,  268 
Ornate  tree  frog,  i,  268 
Ornate  winter  frog,  268 
O'Roke,  E.  C.,  356 
Ortenburgcr,  A.  I.,  46,  112,  113,  135,  138,  166, 

222,  240,  593,  594,  600,  6iT 
Ortcnburger,  R.  D,  135,  138,  166,  593 
Oiton,  G.  L.,  220,  308,  324,  344,  603 
Osborn,  H.,  60 1 
Osama  longtpes,  306 
Over,  W.  H.,  615 
Ovciton,  F.,  127,  142,  212,  214,  226,  347,  356, 

609 
Ovulation,  26-46,  sec  also  Bi ceding  in  each 

species  account 
Owens,  D.  W.,  368,  496,  578,  599,  617 

Pacific  coast  tree  toad,  329;  frog,  429,  520;  hyla, 
329;  tree  frog,  94,  3^9-37>  3^i ;  tree  toad,  329 

Pack,  H.  J.,  167,  255,  617 

Paine,  C.,  281 

Palmer,  E.  L  ,  226,  589 

Palmer,  K.  V.  W.,  226,  333 

Palmer,  M.  E.,  602 

Palmer,  T.  C  ,  613,  614 

Parker,  H.  W.,  569,  583,  5«7 

Parker,  M  V.,  294,  602,  615 

Parks,  H.  B.,  398,  617 

Patch,  C.  L.,  120,  149,  523,  524,  544,  54^,  549, 
552,  587,  59<> 

Paulmier,  F.  C.,  608 

Pawling,  R.  O.,  614 

Pavne,  F.,  409 

Pccquegnat,  W.  E.,  596 

Peeper,  14,  29,  48,  50,  91,  217,  220,  311-16 


Peeper  frog,  250 

Pcnn,  G.  H.,  Jr.,  593 

Pennsylvania,  7,  210,  229,  231,  232,  246,  247, 

248,  253,  500,  504,  613-14 
Perkins,  R.  M.,  594 
Pet  omy scut  gossypinus,  438 
—  polionotus,  434 
Perrier,  E.  and  R.,  104 
Peter,  J.  E.,  606,  607 
Peters,  J.  A.,  600 
Peyton,  J.  H.,  615 
Piatt,  J.,  600,  617 
Pickens,  A.  L.,  216,  281,  614 
Pickerel  frog,  2,  30,  49,  103,  476-80 
Pickering's  frog,  311 
Pickering's  hyla,  311 
Pickering's  hylodes,  311 
Pickering's  tiee  frog,  311 
Pickering's  tree  toad,  511 
Pickwell,  G,  595 
Piersol,  W.  H.,  591 
Pig  frog,  i,  459 
Pilsbry,  H.  A ,  377 
Pine  tree  frog,  319 
Pine  tree  toad,  319 

Pine  woods  tree  frog,  15,  29,  48,  50,  319-24 
Pine  woods  tree  toad,  319 
Pmey  woods  tree  frog,  90 
Pink -snouted  frog,  377 
Piping  tree  frog,  311 
Pirme,  M.  D.,  286,  323,  458,  470 
Plain  meadow  frog,  483-88 
Plains  toad,  164 
Pomsett's  lizard,  397,  398 
Pond  frog,  i,  450 
Pope,  C.  L.,  600 
Pope,   I,   582 
Pope,  P.  H.,  188,  189,  190,  204,  206,  225,  256, 

308,  342,  344,  350,  399,  571,  599,  603,  616 
Pope,  T.  E.  B  ,  620 
Potter,  D.,  605 
Pratt,  H.  S.,  452,  587,  609 
Prtble,  E.  A  ,  250,  251,  310,  476,  525,  549,  550, 

552,  590,  591,  592 
Priest,  Wm.,  12,  13 
Prince  Edward  Island,  533,  592 
Provanchcr,  L.,  592 
Provincial  lists,  590-92 
Pseiidacns,  3,  5,  7,  14,  25,  29,  33,  69,  88,  229- 

80,  232,  238,  242,  243,  329,  335,  361 

—  biachyphona,  7,   16,  21,  22,  35,  45,  70, 
77,  92,  94,  229-34,  235,  272 

—  bnmleyi,  21,  79,  92,  230,  233-36,  248 

—  copn,  272 

—  nignta  darty,  4,  22,  29,  35,  45,  47,  51, 
70,  71,  78,  93,  174,  240-44,  267,  278,  351, 
509,  567 


INDEX 


635 


—  nignta  jcrarium,  4,  7,  17,  21,  29,  35,  45, 
72,  78,  93,  229.  230,  231,  232,  234,  237, 
238,  244-49,  254,  256,  296,  335,  504 

—  nignta  nignta,  3,  4,  16,  21,  31,  32,  54,  79, 
92,  93,  234,  236*40,  251,  263,  434 

—  nignta  septentrtonalis,  4,  21,  31,  35,  45, 
54,  77,  92,  238,  249-53,  267,  551 

—  nignta  tnsenata,  4,  7,  16,  21,  29,  32,  35, 
45,  46,  68,  71,  77,  93,  229,  230,  234,  237, 
242,  244,  248,  253-59,  263,  267,  277 

—  nignta  venncosa,  4,  21,  35,  45,  79,  93,  237, 
258-62 

—  occidentals,  4,  54.  262-65,  279 

—  oculans,  TO,  15,  16,  18,  22,  29,  32,  33, 
35.  37,  48,  50,  69,  77,  92,  235,  263,  264-68, 
278 

—  otnata,  16,  21,  31,  35,  45,  78,  94,  229, 
230,  238,  262,  263,  268-72,  273,  279,  372 

—  strecfan,  21,  35,  46,  54,  70,  78,  92,  94, 
230,  263,  264,  272-80,  373,  509 

Ptetnohyla  fodiens,  577 
Puerto  Rico,  188,  190 
Pusey,  H.  K.,  104 
Putnam,  F.  W.,  126,  127,  60} 
Putnam,  P.  G.,  619 

Quamtance,  C.  W.,  594 

Quebec,   143,   145,   146,  311,  450,  476,  481* 
504,  510,  512,  533,  540,  544,  549,  550,  551, 

592 

Quilhn,  E.  S.,  276,  370,  374 

Quilhn,  R.  D.,  276,  349,  351,  368,  370,  374 

Rain  frog,  2,  14,  341 
Ram  toad,  14,  in 
Rainfall,  n 
Rainy  day  frog,  14 
Ramsey,  E  E  ,  600 

Rana,  3,  18,  19,  20,  26,   30,  33,  53,  96-103, 
159,  210,  217,  257,399-560 

—  aesopus,  1 8 

— •  ateolata  aesopus,  4,  436,  539 

—  aieolata  areolata,  4,  18,  24,  31,  54,  63,  79, 
98,  102,  399-4<>2,  403.  404,  409,  434,  5i6, 
5'9»  537,  538,  539,  540 

—  ateolata  capitot  4,  404,  538,  540 

—  areolata  cnctilosa,  4,  15,  23,  35,  43,  63,  79, 
102,  359,  400,  402-10 

—  awora  atitota,  13,  16,  18,  22,  24,  35,  44, 
54,  61,  78,  99,  100,  TOI,  410-16,  442,  520, 
522,  527 

—  aurora  dtaytonn,  18,  22,  24,  31,  35,  44,  61, 
63*  79»  IOI<  *59,  4i i,  4*5-i8,  442,  475,  522 

—  austncola,  503 

—  bcilandten,  502,  509,  510,  511 

—  boyln  boyln,  4,  13,  16,  18,  22,  24,  30,  31, 


35>  44,  61,  78,  98,  99,  107,  417-25,  429, 
431,  442,  553 

—  boyln  muscosa,  4,  16,  18,  22,  23,  24,  31,  78, 
99,419,420,424-28,555 

—  boyln  sierrae,  3,  4,  16,  18,  22,  24,  33,  35, 
44,  62,  78,  99,  336,  420,  429-34,  555 

—  brachyiephala,  496,  502,  503,  504 

—  cantabrigensis,  see  R.  s.  cantabngensis 

—  capita,  15,  1 8,  24,  30,  35,  43,  49,  63,  75, 
79,  102,  343,  404,  406,  434-39*  47*.  538 

—  cascadae,  4,  24,  31,  35,  44,  62,  78,  100, 
4i  it  439-44t  527*  533 

—  catesbeiana,  4,  12,  13,  18,  19,  20,  24,  26, 
3Q»  3i»  32,  33,  35,  38,  65,  79,  96,  286,  407, 
444-49,  455,  460-66  passim,  469,  475,  536, 
555,  556,  558,  570 

—  clamata,  see  R  clamitans 

—  clamttans,  13,  14,  18,  19,  20,  24,  26,  30, 

32,  33,  35»  39,  49,  66,  79,  98,  286,  296, 
327,  450-54,  457,  458,  463,  469,  502,  519, 
522,  533,  535,  536,  556,  570 

—  fishen,  19,  20,  22,  24,  31,  51,  54,  66,  79, 
ioi»  334,  447,  448,  453-599  473,  SGO,  506, 
5«7,  5i9,  533 

—  fotren,  516,  517 

—  gryho,  13,  14,  15,  18,  19,  20,  24,  30,  32, 

33,  35,  39,  49,  53,  65,  79,  96,  459-66,  467, 
468,  469,  533,  555 

—  halecina,  399,  475,  49^,  500,  508 

—  halecina  betlandtcn,  508,  511,  513 

—  hecf(schen,  4,  17,  19,  20,  24,  31,  49,  53, 
64,  79,  96,  98,  465-72,  555 

—  longipes,  475 

—  montczumac,  18,  475,  506 

—  omiltcmana,  516,  517 

—  onca,  22,  31,  79,  98,   101,  457,  473-77, 
506,  508,  519 

—  omata,  262 
— •  palmipes,  16 

— -  palusftit,  18,  22,  30,  35,  42,  49»  63,  78, 

J«3,  407,  455,  456,  467,  476-8o,  488,  535 
— •  pip/ens  bcrlandtcn,  506-11,  516 

—  pipicns  btachycephala,  511-14 

—  pipiens  burnsi,  3,  4,  5,  483-88,  500 

—  pipiens  fyndtyoht,  5,  488-93,  500 

—  pipiens  onca,  5,  506 

—  pipiens  pipiens,  3,  5,  13,  14,  18,  24,  26, 
35,  42,  43,  49,  63,  64,  79,  101,  102,  306, 
399,  402,  407,  422,  437,  455,  456,  457,  458, 
473-76  passim,  479,   480,  481-84,  485-97 
passim,  498-520,  529,  542,  555 

—  pipiens  sphcnocephala,  5,  7,  13,  18,  24,  26, 
32,  35,  42,  43,  49,  64,  78,  98,   103,  138, 
437»  475,  482,  483,  492-97,  5oo,  5°2,  5<>3» 
509,  510 

—  prctiosa  lutelventns,  7,  22,  30,  35,  42,  62, 
joo,  442,  522,  526,  527-33 


636 


INDEX 


Rana  prcttosa  prettosa,  7,  13,  18,  22,  25,  35,  42, 
43>  54.  62,  78,  100,  411,  440,  442,  451, 
506,520-28,531,532,533 

—  pustulosa,  517 

—  septcntrionalis,  4,  16,  18,  19,  20,  22,  24, 
3i»  35,  43,  53,  66,  79,  98,  451,  455,  533- 
37,  556,  560 

—  sevosat  24,  79,  404,  489,  537-4O 

—  sphenoccphala,  18,  437,  500 

—  sylvattca  cantabngensts,  18,  22,  25,  26,  31, 
33,  35,  42,  54,  78,  99,  54 1,  542,  544*52 

—  sylvattca  lattremtts,  5,   16,   18,  544,  549, 

550,  551 

—  sylvattca  sylvatica,  13,  18,  22,  25,  26,  30, 
32,  35,  42,  43,  61,  62,  78,  99,  269,  437, 
441,  480,  483,  523,  540-44,  546-50  passim 

—  tarahnmarae,  24,  61,  79,  98,  432,  485,  514, 
516,  553-55*  557 

— .  temporana,  442,  475 

—  titncularia,  399 

—  vtrescens,  503,   511,  513,  see  also  R.  pt- 
piens 

—  virestens  areolata,  399 

—  vitescens  biachytephala,  137,  503,  509,  513, 

5M 

—  viresccns  ctrculosa,  399 

—  vtrgatipcs,  4,  14,  15,  19,  20,  22,  24,  30, 
3i,  33,  35,  42,  49,  5i,  53,  67,  79,  96,  45^, 
460,  463,  468,  471,  504,  533,  535,  55<>-<So 

Raney,  E.  C.,  viu,  445,  588,  609,  610 
Range,  5-10,  see  also  each  species  account 
Ranulac,  3,  18,  22,  24,  33,  61-67,  77,  82,  96- 

i»3»  399-56o 
Rathvon,  S.  S.,  613 
Rattler,  i,  220 
Rtagen,  A.  B.,  615 
Red  spotted  toad,  192 
Red-legged  frog,  412 
Redmond,  C.,  607 
Reed,  C.  L.,  563,  566 
Reese,  A.  M.,  613 
Rehn,  J.  A.  G.,  608 
Rhoads,  S.  N,  199,  615 
Rhode  Island,  614 
Riband  snake,  567 
Ribbed  frogs,  76 
Ribbed  toads,  T,  12,  47,  81 
Rice,  F.  L.,  102,  400,  407,  589,  599 
Rich,  J.,  143 

Richardson,  C.  H.,  Jr.,  287,  291,  586 
Richardson,  J.,  550 

Richmond,  N.  D.,  124,  127,  311,  589,  618,  619 
Ricker,  W.  E.,  105,  107,  590 
Ricord's  frog,  32,  95,  373,  377 
Ridgway,  R.,  12,  354,  356,  358,  461,  599 
Rigncy,  C.  C.,  120,  167,  210,  228,  243,  280, 

612 


Ring  frog,  402 

River-swamp  frog,  i,  17,  19,  49,  53,  98,  466- 

Robber  frogs,  3,  10,  81,  82,  97,  280,  368 

Roberts,  H.  L.,  599 

Robertson,  H.  C.,  603 

Rock  frog,  368 

Rocky  Mountain  frog,  416,  533 

Rocky  Mountain  toad,  86,  206-10 

Rodgers,  T.  L.,  105,  329,  330,  458,  473,  475, 
522,  606 

Ross,  G.,  236 

Rough  chorus  frog,  236 

Rugh,  R.,  588 

Ruppell,  E.,  585 

Rutherford,  A.  V.,  372 

Ruthlmg,  P.  D.  R.,  595 

Ruthvcn,  A.  G.,  123,  137,  138,  146,  149,  190, 
214,  215,  216,  255,  301,  302,  336,  514, 
586,  593,  601,  604,  606,  608,  618,  619 

Ryder,  J.  A.,  574,  610 

Sabahan  peeper,  315 

Salamanders,  3 

Salientia,  2;  characters,  2-5;  families,  3,  n,  12; 

number,  4 

Salt  marsh  frog,  i,  150 
Sampson,  L.  V.,  586 
Sanders,  Ottjs,  353 
Saskatchewan,  250,  252,  520,  592 
Savanna  cricket,  i,  14,  217,  264 
Savannah  cricket  frog,  220 
Say,T.,  164,  584 

Scaphiopodidae,  3,  20,  55-57,  80,  81,  82-83 
Scaphiopus,  5,  25,  27,  33,  53,  77,  80,  82-83, 

110-34,  171,  210,  241 
Scaphiopus  couchit,  i,  4,  16,  18,  20,  27,  31, 

34,  39,  56,  57,  76,  77,  83,  110-13,  115,  116, 

132,  135,  169,  170,  373 
-  -  hammondii  bombtjrons,  4,  20,  56,  78,  82, 

83,  116-20,  T2i,  132 

—  hammondn  hammondii,  10,  16,  18,  20,  27, 
3i,  34,  39,  56,  76,  77,  82,  83,  113-17*  H9, 
120,  132,  135 

—  hammondii  intcrmontanus,  4,  20,  56,  82, 
114,  120-23,  149,  333 

—  holbrool(ii  albus,  4,  7,  31,  54,  83,  127-29 

—  holbrootyi  holbroohii,  10,  n,  13,  15,  16,  18, 
20,  26,  27,  33,  34,  39,  46,  48,  53,  55,  76, 
78,  83,  115,  123-27*  130-34,  343 

—  holbroolyt  hurteru,  20,  34,  36,  41,  56,  77, 
83, 130-34*  279,  509,  548 

Schmidt,  F.  J.  W.,  620 

Schmidt,  K.  P.,  188,  212,  215,  216,  224,  358, 
368,  383,  388,  394,  500,  503,  504,  509, 
542,  548,  549,  578,  588,  589,  591,  596,  599, 
600,  602,  605,  611,  614,  616,  617,  620,  621 


INUJbA 


637 


Schneider,  J.  G.,  303 

Schonberger,  C.  F.,  150,  613,  619 

Schoonmaker,  W.,  212 

Sthreber,  J.  C.  D.,  481,  500,  504,  505 

Scraper  frog,  319,  341 

Screaming  frog,  i 

Secondary  sexual  characters,  14-26,  80 

Sensory  line,  sec  Lateral  line 

Seton,  E.  T.,  182,  184,  253,  550,  591 

Shad  frog,  i,  481,  492 

Shannon,  R.  C.,  292,  334 

Shapovalov,  L.,  596 

Shaw,  G.,  4,  444,  584 

Shaw,  J.,  427 

Sherman,  F.,  Jr.,  610 

Sheiwoocl,  W.  L.,  315,  596,  608 

Shoup,  C.  S.,  615 

Shreve,  B.,  381 

Sierra  Mad  re  yellow-legged  frog,  99,  424-28 

Sierra  Nevada  yellow-legged  frog,  99,  429-34 

Stgmodon,  438 

Simpson,  S.  E.  R.,  591 

Size,  10,  n,  20-22,  24,  see  also  each  species 

account 

Skermer,  G.  A.,  377,  379,  380,  598 
Slater,  J.  R.,  46,  106,  107,  150,  411,  412,  416, 

439,  440,  441,  442,  443,  444,  447,  522,  525, 

527,  590»  599,  613,  619 
Slater's  frog,  100,  439-44,  44<> 
Slater's  spotted  frog,  439 
Slatten,  R.,  340,  598 
Slevin,  J.,  154,  196,  294.  329,  3*6,  3<>6,  374. 

410,  419,  422,  424,  427,  447,  455,  457,  458, 

520,   587,   590,   593.   595,   599,   606,   607, 

6i3,  617,  619,  620,  621 
Shpp,  W.,  105,  590,  606,  609 
Small,  H.  B.,  591 
Small -spaded  toad,  146 

Smihsca  bandimi,  352,  sec  also  Hyla  baudinn 
Smith,  A.  G.,  614 
Smith,  A.  L.,  588 

Smith,  C.  C.,  187,  402,  509,  588,  612 
Smith,  D.  S.  C.  H.,  603 
Smith,  E.,  595 
Smith,  H.  M.,  46,  87,   1 1 6,   119,   13°,   *34» 

167,  171,  184,  191,  208,  223,  226,  240,  241, 

243,  244,  257,  353,  4°4,  4»8,  480,  561,  567, 

577,  578,  581,  582,  583,  589,  601,  602,  606, 

611,  616,  617,  621 
Smith,  O.  R.,  1 60,  431 
Smith,  R.  E.,  334,  596 
Smith,  T.  F.,  368,  394,  500,  503,  509,  617 
Smith,  W.  H ,  585,  604,  fii  i 
Snake  frog,  434 
Snyder,  C.  E.,  434 
Snyder,  J.  O.,  121,  122,  606 
Solitary  spadcfoot,  i,  13,  124 


Solitary  toad,  124 

Sollberger,  D.,  231,  561,  567,  617 

Sonmni,  C.  S.,  584 

Sonora  hyla,  i,  360 

Sonora  toad,  172 

Sonora  tree  frog,  360-65 

Sonoran  toad,  167,  184 

Sonoran  tree  toad,  287 

South  Caiolma,  60,  62,  64,  72,  199,  217, 
224,  234,  235,  244,  270,  281,  286,  311, 
341,  360,  434,  447,  465,  481,  495,  504, 
512,  538,  540,  556,  559,  582,  614 

South  Dakota,  116,  180,  206,  344,  353,  355, 
356,  483,  488,  491,  508,  615 

Southeastern  cricket  frog,  217 

Southern  bullfrog,  15,  19,  30,  49,  53,  96,  459 

Southern  California  toad,  85,  156 

Southern  crayfish  frog,  399,  400 

Southern  gopher  frog,  399,  434 

Southern  leopard  frog,  492,  497 

Southern  meadow  frog,  49,  103,  492-98 

Southern  peeper,  315 

Southern  spadefoot,  1 1 1 

Southern  spung  peeper,  315 

Southern  toad,  15,  17,  28,  50,  85,  199 

Southern  tree  frog,  341 

Southern  tree  toad,  341 

Southern  yellow-legged  frog,  424 

Spadefoot,  3,  10,  14,  15,  27,  32,  33,  47,  48, 
5*.  76,  77,  80,  82,  83,  124,  181,  454 

Spadctoot  toad,  15,  28,  51,  85,  124,  167 

Spadefooted  toad,  17,  167 

Spca,  120 

Species  accounts,  104-583 

Speck,  F   G.,  143,  590,  607 

Sphagnum  cricket  fiog,  217 

Sphagnum  frog,  15,  19,  50,  49,  51,  53,  96, 
556-60 

Spiracle,  76 

Spotted  frog,  481,  492,  520 

Spotted  green  frog,  481 

Spotted  toad,  28,  51,  87,  192- 

Spotted  tree  frog,  250 

Spraguc,  J  M.,  119,  60 1 

Spring  fiog,  450,  476,  492 

Spung  peeper,  1,14,  253,  311 

Spimger,  S.,  496,  5*8,  539,  597,  600,  605 

Squinel  frog,  341;  hyla,  341;  tree  frog,  29, 
48,  5",  90,  34I-45J  tree  toad,  341 

Stanton,  K.,  160,  596 

State  lists,  592-620 

StaufTer,  R.  E.  C.,  595 

Stcjncger,  L.,  2,  4,  5,  84,  104,  116,  154-56, 
172,  184,  195,  244,  341,  359,  389,  392,  394, 
397,  399,  402,  454,  45<>,  457,  459,  462,  464, 
473,  487,  506,  509,  512,  583,  588,  589,  593, 
595,  597,  599,  606,  614,  016,  619 


638 


INDEX 


Stewart,  L.,  598 

Stickel,  W.  H.,  597 

Stink  frog,  429 

Stone,  W.,  594,  596,  607,  608,  611,  613,  616 

Storer,  D.  H.,  480,  603 

Storer,  R.,  336,  418,  423 

Storer,  T.  I.,  115,  116,  152,  154,  163,  196, 
288,  291,  294,  333,  336,  411,  416,  417,  418, 
419,  423,  427,  431,  434,  522,  527,  595 

Storm  toad,  124 

Strader,  L.  D.,  619 

Strcckcr,  J.  K.,  Jr.,  112,  113,  130,  131,  133, 
167,  169,  171,  174,  222,  240,  244,  262,  263, 
264,  272-75  passim,  277,  280,  302,  349-52 
passim,  374,  568,  569,  582,  583,  594,  616, 
617 

Strecker's  ornate  chorus  frog,  94,  272 

Street,  J  F.,  607 

Streets,  T.  H.,  620 

Striped  bush  frog,  253 

Striped  chorus  frog,  236,  240 

Striped  tree  frog,  i,  29,  93,  240,  244,  253 

Structural  terms,  adults  (Figs.  1-7),  19,  20,  23, 
24,  25,  80,  88,  97 

Stuart,  L.  C.,  587 

Surface,  H.  A ,  613 

Sutchtf,  R.,  46,  248,  584 

Svihla,  A.,  46,  108,  336,  514,  520,  524,  525, 
526,  532,  619 

Svihla,  R.  D.,  108,  336,  514,  619 

Swamp  bullfrog,  459 

Swamp  chorus  frog,  236 

Swamp  cricket  frog,  14,  77,  92,  93,  236,  244, 
253,  264,  434 

Swamp  frog,  476 

Swamp  tree  frog,  236,  244,  253,  264,  268 

Swamp  tree  toad,  236,  244 

Swamp  whistler,  250 

Swanson,  G.,  487,  491,  605 

Swanson,  P.  L.,  497,  600 

Sweetman,  H.  L.,  347,  604 

Sytrhophus,  81,  94-96,  389-98 

—  campi,  4,  10,  16,  22,  31,  32,  33,  36,  54,  96, 
206,  371,  389-94*  396 

—  gaigac,  4,  7,  22,  31,  54,  95,  371,  394-95 

—  marnocty,  7>  l6>  22,  31,  32,  33,  36,  54, 
95,  37i,  389»  394,  395-98 


Tadpoles,  4^-79 

Tail,  23,  25,  47,  80,  106,  no 

Tail  crests,  47»  75»  ?6 

Tail  musculature,  47,  «*  also  Muscle  seg- 
ments 

Tanner,  V.  M.,  82,  83, 116,  120,  121,  122,  129, 
134,  166,  287,  291,  292,  473,  475,  47$,  5°6, 
519,  588,  617,  618 


Tanner,  W.  W.,  258,  506,  513,  520,  523,  527, 

594,  599,  618,  619 

Taylor,  E.  H.,  87,  103,  173,  174,  184-88  pas- 
stmt  240,  360-63  passim,  365,  367,  383,  384, 
387,  424,  426,  505,  561,  567,  575-78  pas- 
sim, 587,  588,  589,  594,  601,  608,  616,  617, 
618,  621 

Taylor,  W.  P.,  210,  336,  606 

Taylor's  toad,  i,  12,  103,  561-67 

Telford,  H.  S.,  610 

Tennessee,  199,  229,  232,  253,  294,  353,  402, 
481,  540,  615 

Terminology,  see  Structural  terms 

Test,  F.  C  ,  336,  527,  586,  605,  606,  616,  620 

Tevis,  L,  Jr.,  156,329,  621 

Texas,  7,  10,  14,  26,  32,  46,  54,  56,  57,  58, 
60,  63,  64,  72,  73,  74,  79,  111-16  passim, 
124, 130-34  passim,  164,  167,  169-74  passim, 
184-96  passim,  203,  204,  206,  210,  222,  225, 
227,  228,  239-44  passim,  262,  263,  264, 
272-80  passim,  287,  290-92  passim,  298-304 
passim,  306,  311,  319,  341,  342,  344,  347*53 
passim,  360,  366,  367-77  passim,  383-402 
passim,  444,  447,  448,  450,  481,  495,  497, 
500,  502,  503,  504,  508-11  passim,  561-71 
passim,  578,  580,  582,  583,  615-17 

Texas  cliff  frog,  10,  95,  368-77 

Texas  gopher  frog,  102,  399-402 

Texas  narrow  mouth,  578 

Texas  narrow-mouth  toad,  27,  103,  578-83 

Texas  narrow-mouthed  frog,  272 

Texas  ornate  chorus  fiog,  272 

Texas  toad,  164 

Texas  toothless  frog,  578 

Thamnophis  eques,  306 

Thamnopttts  o.  hammondtt,  159 

Thamnophis  o.  hydropMa,  423 

Thamnophis  sauntus  proximus,  204,  206,  567 

Thick-skinned  frog,  i,  419 

Thompson,  C  ,  255,  402,  404,  407,  409,  604 

Thompson,  H.,  255,  409,  525,  527,  529,  53°, 
531,  604,  606,  see  also  Mrs.  II.  T.  Gaigc 

Thompson,  Z,  618 

Thrtc-lmcd  tree  frog,  j,  253 

Three-striped  tree  frog,  253 

Throat,  14-18,  23-25 

Tiger  frog,  476 

Tihcn,  J.  A.,  120,  60 1 

Toad,  3,  10,  14,  32,  33,  47,  77,  81,  83-87,  88, 
122,  245,  434 

Toner,  G.  C.,  478,  550,  589,  590,  592 

Toothless  frog,  i,  571 

Transformation,  75-79 

Trapido,  H.,  145,  503,  504,  549,  551,  618 

Trautman,  M.  B.,  618 

Tree  frog,  13,  33,  52,  82,  87-94,  264 

Tree  toad,  3,  32,  77 


INDEX 


639 


Troost,  G.,  615 

Trowbridge,  A.  H.,  46,  119,  612 

Trowbndge,  M.  S.,  119,  612 

True,  F.  W.,  544 

Tschudi,  J.  J.,  568,  585 

Turner,  L.  M.,  353,  356 

Twitty,  V.  C.,  334,  596 

Uhler,  F.  H.,  356,  359 
Unspotted  meadow  frog,  483 
Upton,  G.  B.,  162 

Utah,  7,  57,  73,  74,  101,  in,  120,  121,  123, 
124,  149,  i65,  166,  167,  171,  197,  206,  209, 

222,  228,  256,  258,  287,  290,  292,  329,  473, 
475,   476,   506,   508,   509,   513,   520,   523, 
617-18 

Utah  frog,  10 1,  473-77 

Valley  cricket  frog,  220 

Van  Deman,  A.  M.,  450 

Van  Deman,  C.  C,,  450 

Van  Deman,  C.  E,  162,  164,  432 

Van  Dcnburgh,  J.,  104,  149,  336,  593,  595» 

599,  606,  608,  613,  617,  620,  621 
Van  Dcital,  W.  R  ,  8,  9,  5°5 
Van  Hymng,  O.  C,  46,  124,  260,  261,  264, 

265,   269,   270,   272,   295,   311,  338,   339, 

341,357,377,  434,  445,  492 
Van  Hymng,  T ,  597 
Van  Vhct,  S.,  511 
Van  Vhet's  frog,  298 
Van  Winkle,  K.,  226,  595,  613,  619 
Varying  tree  toad,  344 
Vegetable  pole,  36 
Ventral  sucker,  75 
Vermont,  500,  504,  618 
Vcrnll,  A.  E.,  604 
Vestal,  E.  H.,  596 
Vigne,  G.  T ,  448,  585 
Vilhers,  C.  G.  S.  de,  104,  587 
Vines,  R.  H.,  400 
Viosca,  Percy,  Jr.,  199,  202,  214,  218,  220,  222, 

223,  234,  268,  294,  296,  298,  319,  352,  356- 
60  passim,  405,  464,  496,  538,  539,  54° 

Viosca's  tree  frog,  i,  91,  294 

Virginia,  14,  54,  73,  74,  M°,  217,  220,  222, 
234,  248,  303,  3«8,  3f>9,  3io>  3",  34i,  445, 
481,  492,  495,  618 

Visher,  S.  S.,  615 

Vitellme  membrane,  34,  35 

Vitellus,  34,  35 

Vladykov,  V.  D.,  145,  592 

Vocal  sacs,  14-18,  23-25 

Vogel,  H.  H.,  Jr.,  60 1,  609 

Voice,  i,  1 2- 1 8,  24-26,  see  also  each  species  ac- 
count 


Von  Bloecker,  J.  C.,  Jr.,  596 
Vorhies,  C.  T.,  368,  376,  377 


Walbaum,  J.  J,  188 

Walker,  C.  F.,  92,  212,  216,  229,  231,  232, 

234,  235,  598,  603,  610,  611,  618,  619 
Wallace,  H.  K.,  261,  271,  324,  471 
Walters,  E.  P.,  234,  602 
Washburn,  F.  L.,  613 
Washington,  7,  44,  46,  55,  62,  105,  106,  108, 

120,  149,  150,  330,  33i,  335,  336,  410-16 

passim,  430,  439-44,  447,  481,  5<>4>  5", 

513,  520-22,  524-27  passim,  619 
Water  frog,  481,  492 
Weber,  C.,  ^54,  359,  547 
Weber,  J.  A  ,  454,  480,  609 
Webs,  25,  80,  97,  1 86 
Weed,  A.  C,  3,  392,  393,  483,  485,  487,  488, 

489,  491,  500,  599,  605 
Wchrle,  Mr.  &  Mrs.  L.  P.,  138,  514 
Welter,  W.  A.,  231,  497,  602 
Werner,  F,  586 
West  cricket  frog,  220 
West  Virginia,  7,  45,  124,  210,  224,  229,  231, 

232,  543,  620 

Western  chorus  frog,  262-64 
Western  cricket,  220 
Western  cricket  frog,  220 
Western  frog,  429,  520-28 
Western  marsh  toad,  150 
Western  narrow-mouthed  frog,  578 
Western  plains  toad,  164 
Western  spadefoot,  113 
Western  spadefoot  toad,  113,  116,  120 
Western  spotted  frog,  100,  520-28 
Western  toad,  164,  167 
Western  tree  frog,  347 
Western  wood  frog,  410,  416 
Wheeler,  R.,  138 
Whistling  frog,  294 
Whistling  tree  frog,  294 
White-jawed  frog,  383 
White-jawed  robber  frog,  383 
White-lipped  frog,  95,  383-88 
Wied,  Prince  Max  zu,  93,  253,  254,  311,  585 
Wicgmann,  A.  F.  A.,  167,  171,  192,  203 
Wiegmann*s  toad,  203-206 
Wiggans,  I.  L.,  160,  176,  178,  596 
Wilcox,  E.  V.,  611 
Wiley,  G.  O.,  302,  486 
Williams,  M.  Y.,  590,  591,  593 
Williams,  W.  J.,  349,  396,  397,  398,  616 
Williamson,  W.  D.,  603 
Wilson,  L.  W.,  224,  543,  618,  620 
Winnipeg,  i,  180,  250 
Winnipeg  toad,  i,  180 


640 


INDEX 


Wisconsin,  253,  355,  359,  489,  502,  504,  511, 

540,  544,  547,  550,  620 
Wood,J.T.,6n 
Wood,  W.  R,  107,  121,  122,  124,  413,  427, 

594,  596,  6:8,  621 
Wood  frog,  i,  30, 32,  99, 329, 540-44 
Woodbury,  A.  M.,  476 
Woodhouse's  toad,  206 
Woods  frog,  540 

Wright,  H.  P.,  407,  408,  409,  600 
Wright,  J.  S.,  188, 189, 190, 383, 616,  618 
Wright's  bullfrog,  465 


Wrights'  tree  frog,  94,  360 
Wyoming,  120,  444,  513,  520,  620 

Yarrow,  H.  C.,  143-46  passim,  154,  475,  476, 
506,  508,  589,  593,  595,  596,  606,  610,  617 
Yellow-legged  frog,  98 
Yellow-throated  green  frog,  450 
Yoder,H.D.,23i,6i4 
Yolk,  see  Vitellus 
Yosemite  Park  toad,  160 
Yosemite  toad,  84,  160 
Youngstrom,  K.  A.,  257, 601